BERGEY'S MANUAL OF DETERMINATIVE BACTERIOLOGY BY ROBERT S. BREED New York State Experiment Station {Cornell University), Geneva, New York E. G. D. MURRAY^ McGill University, Montreal, Province Quebec, Canada A. PARKER HITCHENS University of Pennsylvania, Philadelphia, Pennsylvania Assisted by Sixty Contributors Whose Names and Contributions Appear in the Pages Immediately Following SIXTH EDITION fi/i{irsA^s\\si\ iKVAVTACHE j'At BALTEVIORE THE WILLIAMS & WILKINS COMPANY 1948 First Edition, August, 1923 Second Edition, December, 1925 Third Edition, January, 1930 Fourth Edition, March, 1934 Preprint of pages ix + 79 of Fifth Edition, October, 1938 Fifth Edition, April, 1939 Copyright, 1948 The Williams & Wilkins Company Made in United States of America Published January, 1948 Composed and Printed at thb WAVERLY PRESS, INC. FOS The Williams & Wilkins Compajjy Mt. Royal and Guilford Aves. Baltimore, Md., U. S. A. LIST OF CONTRIBUTORS Allen, O. N. Baldwin, I. L. Barker, H. A. Bengtson, Ida A. Bergey, D. H.f Borman, Earl L. Branham, Sara E. Breed, Robert S. Robert S. and Margaret E. Breed Buchanan, R. E. Burkholder, Walter H. Rhizobium 223 Rhizobium 223 Methanococcus 248 Butyribacterium 380 Methanobacterium 645 Rickeitsiaceae 1083 Bartonellaceae 1100 Methanomonas 179 Mycoplana 191 Thiospira 212 Achromobacter 417 Flavobacterium 427 Dialister 594 Paracolobactrum 460 N eisseriaceae 295 Survey of Classifications 5-38 Rules of Nomenclature 49-64 Nitrobacteriaceae 69 Pseudomonadaceae 62, 171 Chromobacteriurn 231 Methanococcus 248 Pediococcus 249 Sarcina 285 Leptotrichia 264 Corynebacteriaceae 381 Achromobacteriaceae 412 App. 2. Eschericheae 461 Serratia 479 Malleomyces 554 Fusobacterium, 581 and Fusiformis 583 Bacteriaceae 596 Caulobacteriineae 827 Chlamydobacteriales 981 Indexes 1297 How Bacteria Are Named and Identified 39-48 Etymology 64 ff. Myxobacteriales 1005 Pseudomonadaceae 82 and 150 Corynebacterium 381 Erwima 463 Bacterium 638 t Deceased, September, 1937. IV LIST OF CONTRIBUTORS Chapman, Orren D. Klebsiella 457 Donovania 559 Chester, F. D.f Erwinia 463 Bacillus 704 Clise, Eleanore H. App. 3. Micrococcus 252 Pasteurella 546 App. 1. Bacteroides, 575; App. Eubacteriineae, 692; App. Nocardia, 915; App. Streptomyces, 967; and App. Spirochaetales, 1051 Conn, H. J. Nitrobacteriaceae 69 Agrobacterium 227 App. 3. Corynebacterium 407 Alcaligenes 412 Davis, Gordon E. Borrelia 1058 Dienes, Louis Borrelomyceiaceae 1287 Edwards, P. R. Salmonella 492 Evans, Alice C. Parvobacteriaceae 545 Fred, E. B. Rhizobium 223 Hagan, W. A. Parvobacteriaceae 545 Hall, Ivan C. Anaerobic section Micrococcus, 246; Neisseria, 255; Veillonella, 302; Diplococcus, 308; and Strepto- coccus, 328 Hanks, John H. Mycobacterium 876 Harvey, Philip Pasteurella 546 Haynes, Wm. C. Vibrio 192 Spirillum 216 Henrici, A. T.J Caulobacieriineae 827 Nocardia 892 Actinomyces 925 Streptomyces 929 Chlamydobacteriales 981 Hitchens, A. Parker Vibrio 192 Mimeae 595 Editor Supplement I, 1082, II, 1125 and III, 1287 Hitchner, E. R. Aeromonas 101 Hofer, A. W. Azotobacleriaceae 219 Holmes, Francis O. Virales 1200 Hucker, G.J. Micrococcus 235 Gaffkya 283 Streptococcus 312 Leuconostoc 346 Huddleson, I. F. Brucella 560 Johnson, Frank H. Phosphorescent Bacteria 633 Kauffmann, F. Salmonella 492 Kelly, C. D. Acetobacler 179 Bacteroides 564 Noguchia 592 Kirby, Harold App. 1. Rickettsiales 1121 Meyer, K. F. Pasteurella 545 t Deceased, January, 1943. J Deceased, April, 1943. LIST OF CO>n'RIBUTORS Murray, E. G. D. Pederson, C. S. Peshkoff, Michael A. Pittman, Margaret Rahn, Otto Rake, Geoffrey Reed, Guilford B. Rettger, L. F. Robinson, George H.t Roy, T. E. Rustigian, Robert Sherman, J. M. Smith, Frederick Smith, X. R. Speck, M. L. Spray, R. S. Stanier, R. Y. Starkey, R. L. Steinhaus, Edward A. Stuart, C. A. Tobie, W. C. Van Xiel, C. B. Vaughn, Reese Waksman, Selman A. Weiimian, David Welsh, Mark Yale, M. W. ZoBell, Claude E. Neisseriaceae 295 Diplococcus 305 Streptococcus 312 Corynebacteriaceae 381 Parvobacteriaceae 545 Spirochaetales 1051 Leuconostoc 346 Lactobacillus 349 Lepiotrichia 364 Microbacterium 370 Butyribacterium 380 Caryophanales 1002 Hemophilus 584 Characterizations of Groups 64 ff. Chlamydozoaceae 1114 Mycobacteriaceae 875 Lactobacillus 349 Spirochaetales 1051 Bacteroides 564 Proteus 486 Streptococcus 312 Lactobacillus 349 Salmonella 492 Sh igella 525 Bacillus 704 Microbacterium 370 Clostridium 763 Cytophagaceae 1012 Sporocytophagaceae 1048 Xitrobacteriaceae 69 Insect Microbiology 417, 742 ff. Rickettsiaceae 1083 Proteus 486 Chromobacterium 231 Propionibacterium 372 Rhodobacteriineae 838 Bcggiatoaceae 988 Achromatiaceae 997 Acetobacter 179 Actinomycetaceae 892 Streptomycetaceae 929 Bartonellaceae 1100 Pasteurella 546 Escherichia 444 Aerobacter 453 Proteus 486 Desulfovibrio 209 Marine Microbiology 107, 418, 431 ff. t Deceased, October, 1945. PREFACE TO SIXTH EDITION More than the usual amount of time and effort has been given toward making this new edition of Bergey's Manual useful. The volume has been completely revised and is reset in double column format so that each page carries about 20 per cent more type than the pages in the fifth edition. Those who are interested in special groups of bacteria will find something new in the presentation of the relationships in every genus. Because of our rapidly expanding knowledge, changes in the outline classification and text were made necessary. These changes have in every case been made by speciahsts in consultation with the Editorial Board. Every spe- cialist possesses first hand knowledge of the species in the group that he or she has reviewed. Because increasing knowledge has shown the fission fungi to be a larger and more diversified group than previously realized, the number of species described has increased from 1335 in the fifth edition to 1630 in the present edition of the Manual. This number does not cover all of the descriptions found in the literature for, as in all other fields of biology, many of the descriptions are so inadequate that the species described cannot now be identified. Many descriptions are obvious or probable duplications of previous descriptions while still others are based on nothing more substan- tial than the author's belief that he had something new, he having made but little effort to compare his cultures with those found by previous investigators. An indication of the large number of inadequate descrip- tions will be found by referring to the material in the appendixes to the various groups, and to the index where synonyms and incompletely described species are shown in italics. The large number of these poorly described species suggests that there has been much unsatisfactory work done in the field of bacteriological taxonomy. Progress in this inadequately developed field is needed as it would help to clarify the approach to desirable research in many fields of bacteriology. It is believed that both teachers and investigators will find the new Source and Habitat index useful. It is important to know what organisms have been described from any given habitat in determining the identity of a described species or whether a given species is new. The future development of taxonomic work holds several interesting possibilities of increased international cooperation such as between the various National Type Culture Collections and within the International Vm PKEFACE TO SIXTH EDITION Association of Microbiologists. The Trust Funds provided through the generosity of Dr. Bergey before his death have been used in developing the present edition of the Manual and future funds are to be used in the same way under the management of a self-perpetuating Board of Editor-Trustees. We are all under obligation to those who have given so freely of their time and special knowledge in preparing this edition cf the Manual. Moreover the Editor-in-Chief is under special obligation to his wife, Margaret Edson Breed who has carried the burden of the indexing; to Mrs. Eleanore Heist Clise who has given invaluable service in bibliographical research, in proof reading and other ways; and to his secretary, Miss Maude Hogan, who has cared for many difficult manuscripts and a voluminous technical correspondence. Many binomials not previously mentioned in the Manual will be found in the Index of Genus and Species Names. Each new name means that there is a new bibliographic reference in the text. Practically all of the incomplete references of previous editions and all new references have been examined in the original, something that is essential in all accurate taxo- nomic work. The index of names is the most complete list that has ap- peared in the literature and should always be consulted before new genus or species names are proposed. This edition of the Manual has been more than four years in press, thanks to the care that has been taken to make it complete and useful. Through- out, the Editorial Board has had the cooperation and understanding help of the publishers of the book who themselves have been forced to meet and overcome the trying difficulties of the war years. The plan of the present book is such that it will be found useful both to teachers and research workers. Robert S. Breed, Chairman E. G. D. Murray A. Parker Kitchens Board of Editor-Trustee.?. April, 1947. PREFACE OF FIRST EDITION The elaborate system of classification of the bacteria into families, tribes and genera by a Committee on Characterization and Classification of the Society of American Bacteriologists (1917, 1920) has made it very desirable to be able to place in the hands of students a more detailed key for the identification of species than any that is available at present. The valuable book on "Determinative Bacteriologj'^" by Professor F. D. Chester, pub- lished in 1 901 , is now of very little assistance to the student, and all previous classifications are of still less value, especially as earlier systems of classifica- tion were based entirely on morphologic characters. It is hoped that this manual will serve to stimulate efforts to perfect the classification of bacteria, especially by emphasizing the valuable features as well as the weaker points in the new system which the Committee of the Society of American Bacteriologists has promulgated. The Committee does not regard the classification of species offered here as in any sense final, but merel}^ a progress report leading to more satisfactory classification in the future. The Committee desires to express its appreciation and thanks to those members of the society who gave valuable aid in the compilation of material and the classification of certain species. . . . The assistance of all bacteriologists is earnestly solicited in the cori-ection of possible errors in the text; in the collection of descriptions of all bacteria that may have been omitted from the text; in supplying more detailed descriptions of such organisms as are described incompletely ; and in furnish- ing complete descriptions of new organisms that maj' be discovered, or in directing the attention of the Committee to publications of such newly described bacteria. David H. Bergey, Chairman Francis C. Harrison Robert S. Breed Bernard W. Hammer Frank M. Huntoon Committee on Manual. August, 1923. .'^AS^J ■H CONTENTS Introduction 1 Historical Survey of Classifications 5 How Bacteria are Named and Identified 39 Rules of Nomenclature 49 Class Schizomyceies Nageli 65 Order I. Eubacteriales Buchanan 66 Suborder I. Eubacteriineae Breed, Murray and Hitchens 67 Family I. Nitrobacteriaceae Buchanan 69 Tribe I. Niirobacterieae Winslow et al 70 Genus I. Nitrosomoyias Winogradsky 70 Genus II. Nilrosococcus Winogradsky 71 Genus III. Nitrosospira Winogradsky 71 Genus IV. Nitrosocystis Winogradsky 72 Genus V. Nitrosogloea H. Winogradsky 73 Genus VI. Nitrobacter Winogradsky 74 Genus VII. Nitrocystis H. Winogradsky 75 Tribe II. Hydrogenomonadeae Pribram 76 Genus I. Hydrogenomonas Orla-Jensen 76 Tribe III. Thiobacilleae Bergey, Breed and Murray 78 Genus I. Thiobacillus Beijerinck 78 Family II. Pseudomonadaceae Winslow et al 82 Tribe I. Pseudomonadeae Kluyver and Van Niel 82 Genus I. Pseudomonas Migula 82 Genus II. Xanthomonas Dowson 150 Genus III. Methanomonas Orla-Jensen 179 Genus IV. Acetobacter Beijerinck 179 Genus V. Protaminobacter den Dooren de Jong 189 Genus VI. Mycoplana Gray and Thornton 192 Tribe II. Spirilleae Kluyver and Van Niel 192 Genus I. Vibrio Miiller 192 Genus II. Desulfovibrio Kluyver and Van Niel 207 Genus III. Cellvibrio Winogradsky 209 Genus IV. Cellfalcicula Winogradsky 211 Genus V. Thiospira Vislouch 212 Genus VI. Spirillum Ehrenberg 212 Family III. Azotobacteriaceae Bergey, Breed and Murray 219 Genus I. Azotobacter Beijerinck 219 Appendix : Genus A. Azotomonas Stapp 221 Family IV. Rhizobiaceae Conn 223 Genus I. Rhizobium Frank 223 Genus II. Agrobacterium Conn 227 , Genus III. Chromobacterium Bergonzini 231 Family V. Micrococcaceae Pribram 235 Genus I. Micrococcus Cohn 235 Appendix: Genus A. Methanococcus Kluyver and Van Niel 248 Genus B. Pediococctis Balcke 249 XI 7ij Xll CONTENTS Genus II. Gaffkya Trevisan 283 Genus III. Sarcina Goodsir 285 Subgenera: Zymosarcina Smit 285 Methanosarcina Kluy ver and Van Niel 285 Sarcinococcus Breed 285 Sporosarcina Orla-Jensen 285 Family VI. Neisseriaceae Pr^vot 295 Genus I. Neisseria Trevisan 295 Genus II. Veillonella Prevot 302 Family VII. Lactobacteriaceae Orla-Jensen 305 Tribe I. Slreptococceae Trevisan 305 Genus I. Diplococcus Weichselbaum 305 Genus II. Streptococcus Rosenbach 312 Genus III. Leuconostoc Van Tieghem 346 Tribe II. Lactobacilleae Winslow et al 349 Genus I. Lactobacillus Beijerinck 349 Sub-genera: Thermobactcrivii) Orla-Jensen 350 Streptobacterium Orla-Jensen 350 Betabacterium Orla-Jensen 350 Appendix: Genus A. Leptotrichia Trevisan 365 Genus II. Microbacierium Orla-Jensen 370 Genus III. Propionibacterium Orla-Jensen 372 Genus IV. Butyribacterium Barker 379 Family VIII. Corynebacteriaceae Lehmann and Neumann 381 Genus I. Corynebacterium Lehmann and Neumann 381 Genus II. Listeria Pirie 408 Genus III. Erysipelothrix Rosenbach 410 Family IX. Achromobacteriaceae 412 Genus I. Alkaligenes Castellani and Chalmers 412 Genus II. Achromobacter Bergey et al 417 Genus III. Flavobacterium Bergey et al 427 Family X. Enterobacteriaceae Rahn 443 Tribe I. Eschericheae Bergey, Breed and Murray 444 Genus I. Escherichia Castellani and Chalmers 444 Genus II. Aerobacter Beijerinck 453 Genus III. Klebsiella Trevisan 457 Appendix: Genus A. Paracolobactruni Borman, Stuart and Wheeler 459 Tribe II. Erwineae Winslow et al 463 Genus I. Erwinia Winslow et al 463 Tribe III. Serrateae Bergey, Breed and Murray 479 Genus I. Serratia Bizio 3mend. Breed and Breed 479 Tribe IV. Proteae Castellani and Chalmers 486 Genus I. Proteus Hauser 486 Tribe V. Salmonelleae Bergey, Breed and Murray 492 Genus I. Salmonella Lignieres 492 Genus II. Shigella Castellani and Chalmers 535 CONTENTS XIU Family XI. Parvobacteriaceae Rahn 545 Tribe I. Pasteurelleae Castellani and Chalmers 545 Genus I. Pasteurella Trevisan 546 Genus II. Malleomyces Hallier 554 Genus III. Actinobacillus Brumpt 556 Appendix: Genus A. Donovania Anderson et al 558 Tribe II. Brucellcae Bergey, Breed and Murray 560 Genus I. Brucella Meyer and Shaw 560 Tribe III. Bacteroideae Breed, Murray and Kitchens 564 Genus I. Bacteroides Castellani and Chalmers 564 Genus II. Fusohacterium Knorr 581 Appendix : Genus A. Fusiformis Hoelling 583 Tribe IV. Hemophileae Winslow et al 584 Genus I. Hemophilus Winslow et al 584 Genus II. Moraxella Lwoff 590 Genus III. Noguchia Olitsky, Sy vert on and Tyler 592 Genus IV. Dialister Bergey et al 594 Appendix : Tribe Mimeae DeBord 595 Family XII. Bacteriaceae Cohn 596 Genus I. Bacterium Ehrenberg 596 Subgenera: Kurlhia Trevisan 600 Cellulomonas Bergey et al 613 Saccharobacterium Sickles and Shaw 623 Agarbacterium Angst 624 Photobacterium Beijerinck 633 Methanobacterium Kluyver and Van Niel 645 Appendix: Suborder Eubacteriineae: Overlooked species and syn- onyms 692 Family XIII. Bacillaceae Fisher 704 Genus I. Bacillus Cohn 705 Genus II. Clostridium Prazmowski 763 Suborder II. Caulobacteriineae Breed, Murray and Kitchens 828 Family I. Nerskiaceae Kenrici and Johnson 830 Genus I. Nevskia Famintzin 830 Family II. Gallionellaceae Kenrici and Johnson 830 Genus I. Gallionella Ehrenberg 831 Family III. Caulobacteriaceae Kenrici and Johnson 832 Genus I. Caulobacter Kenrici and Johnson 832 Family IV. Siderocapsaceae Pribram , . 833 Genus I. Siderocapsa Molisch 833 Genus II. Sideromonas Cholodny 834 Appendix: Family Pasteuriaceae Laurent 836 Genus I. Pasteuria Metchnikoff 836 Genus II. Blastocaulis Kenrici and Johnson 836 Suborder III. Rhodobacteriineae Breed, Murray and Kitchens 838 Family I. Thiorhodaceae Molisch 841 Genus I. Thiosarcina Winogradsky 842 Genus II. Thiopedia Winogradsky 843 Genus III. Thiocapsa Winogradsky 844 Genus IV. Thiodictyon Winogradsky 845 XIV CONTENTS Genus V. Thiothece Winogradsky 846 Genus VI. Thiocystis Winogradsky 846 Genus VII. Lamprocystis Schroeter 847 Genus VIII. Amoebobacter Winogradsky 848 Genus IX. Thiopolycoccus Winogradsky 860 Genus X. Thiospirillum Winogradsky 850 Genus XI. Rhabdomonas Cohn 853 Genus XII. Rhodothece Molisch 855 Genus XIII. Chromatium Perty 856 Family II. Aihiorhodaceae Molisch 861 Genus I. Rhodopseudomonas Kluyver and Van Niel emend. Van Niel 861 Genus II. Rhodospirillum Molisch 866 Family III. Chlorobacteriaceae Geitler and Pascher 869 Genus I. Chlorobium Nadson 869 Genus II. Pelodictyon Lauterborn 870 Genus III. Clathrochloris Geitler 872 Genus IV. Chlorobacterium Lauterborn 872 Genus V. Chlorochromatium Lauterborn 873 Genus VI. Cylindrogloea Perfiliew 873 Order II. Actinomycetales Buchanan 875 Family I. Mycobacteriaceae Chester 875 Genus I. Mycobacterium Lehmann and Neumann 876 Family II. Actinomyceiaceae Buchanan 892 Genus I. Nocardia Trevisan 892 Genus II. Actinomyces Harz 925 Family III. Streptomycetaceae Waksman and Henrici 929 Genus I. Streptomyces Waksman and Henrici 929 Genus II. Micromonospora Prskov 978 Order III. Chlamydobacteriales Buchanan 981 Family I. Chlamydobacteriaceae Migula 981 Genus I. Sphaerotilus Kiitzing 982 Genus II. Clonothrix Roze 983 Genus III. Leptothrix Ktitzing 983 Family II. Crenothricaceae Hansgirg 987 Genus I. Crenothrix Cohn 987 Family III. Beggiatoaceae Migula 988 Genus I. Thiothrix Winogradsky 988 Genus II. Beggiatoa Trevisan 990 Genus III. Thiospirillopsis Uphof 993 Genus IV. Thioploca Lauterborn 993 Appendix: Family Achromatiaceae Massart 997 Genus I. Achromaiium Schewiakoff 997 Genus II. Thiovulum Hinze 999 Genus III. Macromonas Utermohl and Koppe 1000 Appendix : Order Caryophanales Peshkoff 1002 Family I. Pontothricaceae Peshkoff 1002 Genus I. Pontothrix Nadson and Krassilnikow 1002 Family II. Arthromitaceae Peshkoff 1002 Genus I. Arthromitus Leidy 1002 Genus II. Coleomitus Duboscq and Grass^ 1003 CONTENTS XV Family III. Oscillospiraceae Peshkoff 1003 Genus I. Oscillospira Chatton and Perard 1004 Family IV. Caryophanaceae Peshkoff 1004 Genus I. Caryophanon Peshkoff 1004 Order IV. Myxobacteriales Jahn 1005 Family I. Cytophagaceae Stanier 1012 Genus I. Cytophaga Stanier 1012 Family II. Archangiaceae Jahn 1017 Genus I. Archangium Jahn 1017 Genus II. Stelangium Jahn 1020 Family III. Sorangiaceae Jahn 1021 Genus I. Sorangium Jahn 1021 Familj' IV. Polyangiaceae Jahn 1025 Genus I. Polyangium Jahn 1025 Genus II. Synangium Jahn 1032 Genus III. Melittangium Jahn 1033 Genus IV. Podangium Jahn 1034 Genus V. Chondromyces Berkeley and Curtis 1036 Family V. Mycococcaceae Jahn 1040 Genus I. M yxococcus Thaxter 1040 Genus II. Chrondrococcus Jahn 1044 Genus III. Angiococcus Jahn 1047 Genus IV. Sporocyfophaga Stanier 1048 Order V. Spirochaetales Buchanan 1051 Family I. Spirochaetaceae Swellengrebel 1051 Genus I. Spirochaeta Ehrenberg 1051 Genus II. Saprospira Gross 1054 Genus III. Cristispira Gross 1055 Family II. Treponemataceae Schaudinn 1058 Genus I. Borrelia Swellengrebel 1058 Genus II. Treponema Swellengrebel 1071 Genus III. Leptospira Noguchi 1076 Supplement 1 1081 Order Rickettsiales Gieszcykiewicz 1083 Family I. Rickettsiaceae Pinkerton 1083 Genus I. Rickettsia da Rocha-Lima 1084 Genus II. Coxiella Bengtson 1092 Genus III. Cowdria Bengtson 1094 Family II. Bartonellaceae Gieszcykiewicz 1100 Genus I. Bartonella Strong, Tyzzer and Sellards 1100 Genus II. Haemohartonella Tyzzer and Weinman 1102 Genus III. Grahamella Brumpt 1109 Genus IV. Eperythrozoon Schilling 1111 Family III. Chlamydozoaceae Moshkovskj^ 1114 Genus I. Chlamydozodn Halberstaedter and von Prowazek 1114 Genus II. Miyagawanella Brumpt 1115 Genus III. Colesiota Rake 1119 Appendix : Genus A. Caryococcus Dangeard 1121 Genus B. Drepanospira Petschenko 1122 Genus C. Holospora Haffkine 1122 XVI CONTENTS Supplement II 1125 Order Virales Breed, Murray and Kitchens , 1128 Suborder 1. Phagineae Holmes 1128 Family I. Phagaceae Holmes 1128 Genus I. Phagus Holmes 1128 Suborder II. Phytophagineae Holmes 1145 Family I. Chlorogenaceae Holmes 1145 Genus I. Chlorogenus Holmes 1146 Genus II. Carpophthora McKinney 1151 Genus III. Morsus Holmes 1153 Genus IV. Aureogenus Black 1154 Genus V. Galla Holmes 1157 Genus VI. Fractilinae McKinney 1159 Family II. Marmoraceae Holmes 1163 Genus I. Marmor Holmes 1 163 Genus II. Acrogenus Holmes 1202 Genus III. Corium Holmes 1203 Genus IV. Nanus Holmes 1206 Genus V. Rimocortius Milbrath and Zeller 1208 Genus VI. Adelonosus Brierley and Smith 1211 Family III. Annulaceae Holmes 1212 Genus I. Annulus Holmes 1212 Family IV. Rugaceae Holmes 1218 Genus I. Ruga Holmes 1218 Family V. Savoiaceae Holmes 1221 Genus I. Savoia Holmes 1221 Family VI. Lethaceae Holmes 1223 Genus I. Lethum Holmes 1223 Suborder III. Zoophagineae Holmes 1225 Family I. Borrelinaceae Holmes 1225 Genus I. Borrelina Holmes 1225 Genus II. Moralor Holmes 1227 Family II. Borreliotaceae Holmes 1229 Genus I. Borreliota Goodpasture 1229 Genus II. Briareus Holmes 1233 Genus III. Scelus Holmes 1234 Genus IV. Hostis Holmes 1239 Genus V. Moliler Holmes 1241 Family III. Erronaceae Holmes 1248 Genus I. Erro Holmes 1248 Genus II. Legio Holmes 1257 Genus III. Formido Holmes 1263 Family IV. Charonaceae Holmes 1265 Genus I. Charon Holmes 1265 Genus II. Tarpeia Holmes 1268 Genus III. Tortor Holmes 1275 Family V. Trifuriaceae Holmes 1282 Genus I. Trifur Holmes 1282 Family VI. Rabulaceae Holmes 1284 Genus I. Rabula Holmes 1284 Supplement III 1287 Family Borrelomycetaceae Turner 1291 Genus I. Ascococcus Borrel et al 1291 INTRODUCTION Suggestions for the Use of the "Manual in Classifying Unknown Organisms No organism can be classified before we have determined, through de- tailed study, its morphological, cultural, physiological and pathogenic characters. The characters used in the keys to orders, families and genera may ordi- narily be determined by the use of a dozen or more of the procedures de- scribed in the ^vlanual of Pure Culture Study issued by the Committee on Bacteriological Technic (H. J. Conn, Chairman, Geneva, New York) of the Society of American Bacteriologists. j\Iore complete examinations must be made as indicated in the iNIanual of Pure Culture Study, and in the Descriptive Charts which accompany this Manual where it is desired to identify individual species. These tests must be made if bacteria are to be accurately identified and described. It is urged that beginning students be taught the technics necessary for the identification of species in the hope that the taxonomic work of the future may be placed on a more satisfactory basis. After a complete study of the characters of the organism has been made, turn to page 65 and ascertain first in which order the organism belongs. When the order and suborder (if necessary) have been ascer- tained, turn to the page of the IManual on which the key to that order or suborder is given. In this key ascertain the family or subfamily to which the organism belongs. When the family or subfamily has been decided on, again refer to the page of the Manual on which the key to that familj^ or subfamily is given. In this key ascertain the tribe to which the organism belongs. When the tribe has been decided on, again find the page of the Manual on which the key to the tribe is given. In this key ascertain the genus to which the organism belongs. When the genus has been decided on, again refer to the page of the Manual on which the key to that genus is given. In this key, trace out the species under investigation. For example, if one wishes to trace a short, peritrichous, Gram-negative, non-spore-forming rod that grows well on ordinary culture media at 37°C, fermenting glucose and lactose with production of acid and gas, not liquefying gelatin, producing no pigment on any culture medium, with negative reaction for acetylmethylcarbinol, producing indole and reducing nitrates, consult the key to the orders on page 65. 1 2 SUGGESTIONS FOR USE OF MANUAL In this key examine A. Cells rigid, not flexuous. This indicates our or- ganism as its cells remain constant in form. We next examine 1. Cells single, in chains or masses. Not branching and mycelial in character. Not arranged in filaments. Not acid fast. As the organism in question occurs as single cells or at most as short chains and is not acid fast, this indicates that it belongs to the Order Eubacteriales. We now examine a. Do not possess -photo synthetic pigments. Cells do not contain free sulfur. As our organism is unpigmented and the cells do not contain free sulfur, this indicates that our organism belongs to the Sub-order Eubacteriineae. We note that the key to this suborder is on page 67. We next attempt to ascertain the family to which the organism belongs by tracing it through the key to the familiesof the Sub-order Eubacteriineae, p. 67. I. No endospores indicates our organism. We proceed to A. Can develop on inorganic media. As the organism cannot grow without organic carbon, we turn to B. Cannot develop on inorganic media. This corresponds with the physiology of our organism; so we turn to 1. Polar flagellate, etc. As our organism is peritrichous, we proceed to 2. Large oval, pleomorphic cells sometimes almost yeast-like in appearance. Free living in soil. Fix free nitrogen. As this does not correspond with the morphology or physiology of our organism, we next examine 3. Peritrichous or non-motile rods, and cocci. This corresponds with the characteristics of our organism. We turn to a. Heterotrophicrodswhichmaynot require organic nitrogen for growth. Usually motile with one to six or moreflagella. Usually form nodules or tubercles on roots of plants, or shoio violet chromogenesis. This again does not indicate our oi canism ; so we turn next to aa. Heter- otrophic rods or cocci iDhich utilize organic nitrogen and usually carbohydrates. As our rod-shaped organism prefers a medium containing organic nitrogen, we proceed to b. Spherical cells in masses, tetrads and packets. This does not correspond to the morphology of our organism, and we now proceed to bb. Spherical cells ivhich grow in pairs and chains; and rods. This includes our rod-shaped organism; so we turn to c. Gram-positive cocci and rods. Non-motile. Since these are not the characteristics of our organism, we turn to cc. Gram-negative rods. When motile, from four to many peritrichous flagella. Our organism is Gram-negative and peritrichous; so we proceed to d. Grow well on ordinary media containing peptone. Aerobic to factdtative anaerobic. This corresponds with the characteristics of the organism we have studied; so we turn next to e. Gram-negative, straight rods which ferment SUGGESTIONS FOR USE OF MANUAL 3 sugars with the formation of organic acids. This again corresponds with our organism. We turn next to f. Produce little or no acid from litmus milk. This does not correspond with the characters we have determined for our organism. We proceed to ff. Produce CO2 and frequently visible gas (CO2 + H2) from glucose. Reduce nitrates, etc. Our organism produces visible gas from glucose and reduces nitrates. This indicates that it belongs to Family X. Enterohacteriaceae, p. 443. This appears to fit our unknown organism. We now refer to page 443 on which the key to the Family Enterohacteriaceae is found. In this key we ascertain the Tribe to which our organism belongs. 1. Ferment glu- cose and lactose with the formation of acid and visible gas. Usually do not liquefy gelatin. Tribe I. Eschericheae. This corresponds with the characters exhibited by our organism. We refer to the key for Tribe I. Eschericheae on the same page. 1. Methyl red test positive. Voges-Proskauer test negative. Salts of citric acid may or may not be used as sole source of carbon. Genus I. Escherichia, p. 444. This description appears to correspond with that of our unknown or- ganism. We find the key to the species of Genus Escherichia follows the key to the Tribe Eschericheae. On tracing our organism in this key we find that it corresponds to Escherichia coli. A brief description of this or- ganism is found on the same page. * In the use of keys for identifying bacteria, the student is confronted with two difficulties, both based primarily on lack of knowledge and ex- perience. The first is insufficient knowledge concerning the morphology, physiology, possible pathogenicity and habitat of the microorganisms that are to be identified. This may be due to careless observations or to poor training in the special techniques that must be used in determining the identity of a given bacterium. The second difficulty in the use of a key comes from inexperience in the use of technical terms; that is, the student may not thoroughly understand the meaning of the statement in the key and so cannot follow a route through the key with certainty. For example in the keys used here, the student must know the difference (1) between chains of cells which are composed of dividing cells which do not separate at once, and (2) filaments which are composed of dividing cells which remain more permanently to- gether and are normally flattened against each other on adjacent sides. They may show some differentiation into hold fast cells and reproductive cells (conidia), (3) Both chains of cells and filaments are to be distinguished from the mycelial threads found in Actinomycetaceae. These are unseptate and branching with a true branching. * Condensed and paraphrased from Hitchcock's Descriptive Systematic Botanj^, New York, 1935. 4 SUGGESTIONS FOR USE OF MANUAL The student should be warned not to take descriptions in the Manual too literally or too rigidly. Descriptions are usually drawn to represent average findings. Especially among bacteria, characters such as sugar fermentations, gelatin liquefaction, presence or absence of flagella and other things will vary. Sometimes these variations are due to slight, possibly unrecognized variations in the techniques used in determining these characters. Real knowledge of the characteristics of species may also be very incomplete. This is true not only of the physiological activities of these microorganisms; but also in regard to such detectable structural features as the number and position of flagella. Dark field movies of motile cells and photographs taken with the recently developed electron micro- scope are revealing new and heretofore unsuspected facts regarding struc- tural features. Source and habitat data are frequently helpful in aiding the student to recognize species of bacteria and may indicate that the pathogenicity of the culture in question may need to be tried on some specific animal or plant. By habitat is meant the kind of a place in which the organism normally grows; by source, the particular material and place from which the culture was obtained. This source may or may not indicate the natural habitat. The source of cultures is invariably more limited in scope than the habitat as bacteria normally occur wherever their particular habitat may be found in a world wide distribution. The student is also reminded that it is impracticable to note all exceptions in keys. Bacteria like other living things are classified according to a combination of characters, not according to some single character, and ex- ceptions to the characters noted in the keys will occur in nature. These may not be known to or may have been overlooked by the author of the key. On the other hand, the importance of such exceptions should not be over- emphasized and the student would do well to use the key as if there were no exceptions. HISTORICAL SURVEY OF CLASSIFICATIONS OF BACTERIA, WITH E^IPHASIS ON OUTLINES PROPOSED SINCE 1923* There have been numerous attempts to arrange the species of bacteria in natural systems of classification. The first simple system of JMiiller (Ver- mium terrestrium et fiuviatilium, 1773) which he developed further a few- years later (Animalcula infusoria fluviatilia et marina, 1786) listed but two genera {Vibrio and Monas) that included organisms that would today probably be accepted as bacteria. Polyangium Link (^lag. d. Ges. Natur- forsch. Freunde zu Berlin, 3, 1809, 42) is apparently the oldest of the generic terms retained in its original meaning for a bacterial genus while Serratia Bizio (Biblioteca italiana o sia giornale de lettera, scienze ed arti, SO, 1823, 288) was proposed only fourteen years later. Systems of classification developed after 1773 are given in complete outline form in the first edition of the ^Manual (1923) and this section of the IManual was reprinted without material change in the second (1925) and third (1930) editions. While it is not felt to be necessary to repeat these outlines in their entirety, sufficient reference is made below to permit the student to trace the origin of generic terms that are no longer commonly found in classification outlines. No attempt has been made to include reference to other little used generic terms except as the}- appear as syn- onyms in the descriptive portion of the ^Manual. For the origin of generic terms proposed before 1925, see Enlows (The Generic Names of Bacteria, Bui. No. 121, Hygienic Laborator}-, Washington, D. C, 1920) and Bu- chanan (General Sj-stematic Bacteriology, Baltimore, 1925). Bory St. Vincent (IMicroscopiques, Dictionnaire classique d'histoire naturelle, 10, 1826, 533) introduced the generic terms Spirilina, Melanella, Lactrimatoria and Pupella and accepted Vibrio for microorganisms, some of which must have been bacteria. None of these terms, except Vibrio, are in current use for bacterial groups. Three of the terms accepted or proposed by Ehrenberg (Die Infusions- tierchen als volkommene Organismen, Leipzig, 1838); nameh^, Vibrio, Spirillum and Spirochaeta, are still used. The generic term Bacterium proposed first b}- Ehrenberg in 1828 (SjTiibolae Physicae seu Icones et Descriptiones Animalium Evertebratorum Separasitis Insectis quae ex Itinere per Africam Borealem et Asiam Occidentalem, IV. Evertebrata, Berlin) to include but a single species Bacterium triloculare from an oasis * Contributed by Prof. R. S. Breed, New York State Experiment Station, Geneva, New York, July, 1938; revised, September, 1943. b MANUAL OF DETERMINATIVE BACTERIOLOGY in North Africa, has had a varied history because this type species (mono- typy) is no longer identifiable. It was reintroduced into the classification employed in the fifth edition of the Manual to cover species of non-spore- forming rods whose positions in the outline given in the Manual have not yet been satisfactorily determined (Breed and Conn, Jour. Bact., 31, 1936, 517) and is used in the present edition with the same meaning. The term Spirodiscus was applied by Ehrenberg to a single organism that he found in a mountain stream. It has never been reidentified and subsequent authors have discarded this term. Two new generic terms {Metallacter, Sporonema) were introduced by Perty (Zur Kenntniss kleinster Lebensformen, 1852). Neither Metallacter nor Sporonema is in common use at the present time. Davaine (Dictionaire encyclop. des sciences med., Art. bacteries, 1868) introduced one new generic term, Bacteridium, for straight motionless rods like the anthrax bacillus. The generic terms employed by Cohn in his first classification (Unter- suchungen iiber Bakterien. I. Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 146) are all in current use. Only one {Bacillus) was new. Other generic terms were introduced into his second paper (Untersuchungen tiber Bakterien. II. ibid., 1, Heft 3, 1875, 141) which contained his more complete classification. For various reasons, six of these, Merismopedia, Clathrocystis, Ascococcus, Myconostoc, Cladothrix and Streptothrix are not found in recent bacteriological classifications. Mangin (Les Bacteries, Paris, 1878) recognized three subgenera of the genus Monas, the first of which Rhabdomonas Cohn, 1875 is still used as a generic term, while the other two, Ophidomonas Ehrenberg, 1838 and Spiro- monas Perty, 1852 have been dropped. The bacterial species that had been placed in the genus Clathrocystis by Cohn (1875) were separated and placed in a new genus Cohnia by Winter (Die Pilze in Rabenhorst's Kryptogamen Flora, 1880), and this name is also used by Burrill (The Bacteria, Springfield, 111., 1882). Because this name had previously been proposed for a genus of lilies, it was soon dropped. Zopf (Die Spaltpilze, Leipzig, 1883) accepts Phragmidioihrix, a generic name suggested by Engler in 1882 for a single species found on the body of a crustacean {Gammarus locusta). Later authors generally either merge this genus with Crenothrix Cohn or disregard it because of the indefinite description of the one species included in it. Baumgarten (Lehrbuch der pathologischen Mykologie, Braunschweig, 1890) following Hueppe accepts the term, Spirulina, for a genus of pleo- morphic bacteria, disregarding the previous use of the term by algologists. The generic terms found in Migula's first outline (Bakterienkunde fiir SURVEY OF CLASSIFICATIONS OP BACTERIA 7 Landwirte, Berlin, 1890) were those in conventional use at the time and many of them continue in use. Two new terms were introduced for motile types in his second outline (Ai'b. Bact. Inst. Karlsruhe, 1, 1894, 235) and are also found in his later outlines (Engler and Prantl, Die natiirlichen Pflanzenfamilien, 1, la, 1895, 29, and System der Bakterien, 1, 1897, 46, and 2, 1900, 269 and 275) which have not been generally felt to be necessary by subsequent authors. These are Planococcus and Planosarcina. Spiro- soma introduced by Migula in 1894 and Rhabochromatium Winogradsky ac- cepted by Migula in 1900 are like^\ise no longer generally used. Newskia (original spelling Nevskia Famintzen, Bull. Acad. Imp. Sci., St. Petersburg, 34 (N.S. 2), 1892, 484) has recently been revived by Henrici and Johnson (Jour. Bact., 29, 1935, 3 and 30, 1935, 83). The generic term Microspira Schroeter, accepted by Migula in 1894, is still frequently accepted in place of Vibrio as many regard it as having a better status than the later term. The term Pseudomonas was first proposed for polar flagellate bacteria by Migula in his 1894 outline with reference to but a single species, Pseudo- monas violacea, an organism which later investigators have shown to be peritrichous (Cruess-Callaghan and Gorman, Sci. Proc. Roy. Dublin Soc, 21, 1935, 213). Pseudomonas was repeated in the 1895 outline with descriptions of Pseudomonas pyocyanea and other species. Later authors have generally accepted the term Pseudomonas as valid. Fischer (Jahrb. f. mssensch. Bot., Berlin, 27, 1895, 1) introduced a logical outline classification in which he proposed various new terms which have never come into general use. These are Paracloster, Paraplectrum, Arthrobader, Bactrinium, Clostrinium, Plectrinium, Arthrobactrinium, Bactrillum, Clostrillium, Plectrillium, Arthrobactrillium, Badridium, Plec- tridium, Dipledridium, and Arthrobadridium. In his modified classifica- tion (Vorlesungen iiber Bakterien, 1897), he also accepts Pediococcus Balcke, a term that has fallen into disuse except in the brewing indus- try. In the conservative classification proposed bj'' Lehmann and Neumann (Atlas und Grundriss der Bakteriologie, 2 vols., 1896, Miinchen), inter- nationally accepted rules of nomenclature were followed. All of the generic terms employed by them are still in current use, their most important contribution being their acceptance of the suggestion that the genus Bacillus be separated from the genus Baderium on the basis of endospore formation by the rods included in Bacillus. Two new genera were proposed {Corynebaderium and Mycobacterium) that have been generally accepted by later workers. No new generic terms are proposed by Chester either in his preliminarj'^ reports (Delaware College of Agriculture, 9th Ann. Kept., 1897, 53 and 62; 8 MANUAL OF DETERMINATIVE BACTERIOLOGY 11th Ann. Rept., 1899, 36), or in his complete outhne (Manual Determ. Bact., 1901). Almost all of the generic terms found in his outlines are still in current use. The term Aplanohacter suggested by Erwin F. Smith (Bacteria in Rela- tion to Plant Diseases, 1, 1905, 171, Washington) was accepted by certain American phytopathologists for a time but has never come into general use. Because other differences between the non-chromogenic and chromogenic micrococci are unimportant, two generic terms, Albococcus and Aurococcus, suggested by the Winslows (Science, 21, 1905, 669; Systematic Relation- ships of the Coccaceae, New York, 1908) have not come into general use. They also suggested Rhodococcus to include Rhodococcus roseus and R.fulvus apparently without realizing that Zopf (Ber. d. deutsch. bot. Gesellsch. Berlin, 9, 1891, 28) had previously used the same term for Rhodococcus erythromyxa and R. rhodochrous. Hansgirg (Engler and Prantl, Die natiirhchen Pflanzenfamilien, 1, la, 1895, 52) had also used it previously to designate a sub-genus of the green algae, and later Alolisch (Die Purpur- bakterien, Jena, 1907, 20) used Rhodococcus for a genus of the purple bacteria to include Rhodococcus capsulatus. In his complete outline of the classification of bacteria presented in 1909, Orla-Jensen (Cent. f. Bakt., II Abt., 22, 1909, 305) introduced many new generic terms in an effort to create a nomenclature that appeared to him to express the natural relationships of bacteria more satisfactorily than names previously suggested had done. Thus he used the suffixes coccus and sarcina for spherical bacteria and monas for all genera known to be lopho- trichous or so related to these types that they were regarded as essentially lophotrichous in nature. In the same way the suffix bacterium was used for genera of non-spore-forming rods that were regarded as essentially peritrichous in nature, and the suffix bacillus for similar spore-forming rods. As, however, subsequent investigators have (1) accepted the priority rule, (2) felt that it was impossible to recognize the type of motihty found in the ancestry of truly non-motile groups, or (3) felt that other characters were more fundamental than those selected by Orla-Jensen, many of these terms have not been generally used by later workers. Among the little used terms suggested or accepted by Orla-Jensen are: Acetimonas, Nitromonas, Azotomonas, Rhizomonas, Corynemonas, Myco- monas, Sulfomonas, Thiomonas, Thiococcus, Rhodomonas, Rhododictyon, Amoebomonas, Rhodopolycoccus, Rhodosarcina, Spirophyllum, Denitro- monas, Liquidomonas, Liquidovibrio , Liquidococcus, Solidococcus, Solido- vibrio, Sporosarcina, Denitrobacterium, Caseobacterium, Liquidobacterium, Urobacillus, Butyribacillus, Pectobacillus, Cellulobacillus, Putribacillus and Botulobacillus. SURVEY OF CLASSIFICATIONS OF BACTERIA 9 While Nitromonas is not new, it is redefined as a synonym of Nitrohader Winogradsky (Arch. Sci. Biol. St. Petersburg, 1, 1892, 87), rather than as a synonym of Nitromonas Winogradsky (Ann. Inst. Past., 3, 1890, 258). Spirophyllum is from Ellis (Cent. f. Bakt., II Abt., 19, 1907, 507). In a later monograph on The Lactic Acid Bacteria (Mem. d. Acad. Roy. Sci. et Lettres de Danemark, Sect. Sci., 8 Ser., 5, 1919, No. 2) Orla-Jensen proposes the following additional generic terms: Betacocciis,Betahacteriuni, Streptobacterium, Thermohacteriiim and Microhacterium. The term Tctra- coccus is introduced with a meaning different from that given the term previously by v. Klecki (Cent. f. Bakt., 15, 1894, 354). Buchanan prepared an outline classification in 1916 (Jour. Bact., 1, 1916, 591 ; 2, 1917, 155, 347, 603; 3, 1918, 27, 175, 301, 403, 461, 591) which was utilized in part by the group of which he was a member (Winslow, Broadhurst, Buchanan, Krumwiede and Smith) in their preliminary Report to the Society of American Bacteriologists (Jour. Bact., £, 1917, 552) and in the final report by Winslow, Broadhurst, Buchanan, Krumwiede, Rogers and Smith (Jour. Bact., 5, 1920, 191). Although prepared earlier, some parts of the Buchanan outline were not published until after the first Winslow et al. report. As these reports formed the most important basis for the classification used in the first edi- tion of the Manual, it is natural that the generic terms utilized are, in general, the same as those used in the Manual. Generic and subgeneric terms included by Buchanan that are not used in the present edition of the Manual are: Paraspirillum Dobell (Arch. f. Protistenk., 21^, 1911, 97), Eubacillus Hansgirg (Osterr. Bot. Ztschr., 38, 1888, 264; not Eubacillus Dangeard, Le Botaniste, 2, 1891, 151) and Meta- bacterium Chatton and Perard (Comp. rend. Soc. Biol., Paris, 65, 1913, 1232). Siderocapsa Molisch (Ann. Jard. Bot. Buitenzorg, Ser. 2, Supp. 3, 1909, 29) used by Buchanan but dropped by Winslow et al (Jour. Bact., 2, 1917, 549) does not appear in the Manual classification outline until the present (6th) edition. The term Mycoderma recognized both })y Buchanan (Jour. Bact., 3, 1918, 45) and in the preliminary Winslow et al. report (Jour. Bact., 2, 1917, 551) was replaced by the later and more valid term Acetobacter in the final report by WinsloAv et al. (Jour. Bact., 5, 1920, 201). Pfeifferella Buchanan (Jour. Bact., 3, 1918, 54) which is used in the three outline classifications under discussion and also in the first, second and third editions of the ^Ianual, appeared in the literature through a clerical error (Buchanan, General Systematic Bacteriology, 1925, 420) . It was combined in the fourth edition of the Manual with the genus Actinobacillns under the latter name. Nocardia Trevisan (1889) used by Buchanan and in the preliminary report by Winslow et al. (1917) was merged with Actinomyces 10 MANUAL OF DETERMINATIVE BACTERIOLOGY Harz (Jahresber. Miinchen. Thierarzneisch. for 1877-78, 125) in the final report by Winslow et al. Erythrohacillus Fortineau (Compt. rend. Soc. Biol. Paris, 58, 1905, 104) is used by Winslow et al. (1920) but was not accepted in the first and following editions of the IVIanual as it is a synonym of the older Serratia Bizio (1823). Moreover, the species which must be accepted as type for the genus {Erythrohacillus pyosepticus Fortineau (monotypy)) is a species which has been reported by Breed (Manual, 3rd ed., 1930, 117) to be a variant of the older Serratia marcescens. One of the most unsatisfactory portions of recent classifications, such as those outlined by Buchanan (1917-18) and by Winslow et al. (1917), is the treatment given the organisms of the coliform-dysentery-typhoid group in that the term Bacterium is retained for these as suggested by Orla- Jensen (1909). A strict limitation of Bacterium to this group gives it a still different meaning from that which it had had in previous and current classifications, and makes it necessary to find some other place for many other species of Gram-negative, non-spore-forming rods, some of which are well known and well described. The relationships of these miscellaneous species to other non-spore-forming rods is frequently poorly understood. In some cases, further study will probably show that thej'' should be placed in well known and currently recognized genera. In others, further study will probably show that some of these species of non-spore-forming rods should be grouped in new genera. Winslow et al. (1920) recognized this situation and broadened their defini- tion of Bacterium thereby placing such w^ll known species as are included in the colon-dysentery-typhoid group with other species of non-spore-form- ing rods of quite a different character. For this reason, partial use was made in the first edition of the Manual of the numerous generic terms newly proposed by Castellani and Chalmers (Manual of Tropical Medicine, 3rd ed., 1919). Thus the following new terms were introduced: Alcaligenes, Salmonella, Escherichia and Encapsulatus; and the earlier terais Aerohacter Beijerinck (1900) and Eberthella Buchanan (1918). Later it was found that Encapsulatus was a synon3^m of Klebsiella Trevisan (1887), so that the latter term was accepted in the second and subsequent editions of the Manual. Shigella Castellani and Chalmers was recognized as distinct from Eberthella in the third and subsequent editions. Many of the new terms suggested by Castellani and Chalmers were, however, synonyms of earlier valid terms or have not been considered necessary, and so they have not come into general use. These are Nigro- coccus, Graciloides, Cloaca, Eberthus, Dysenteroides, Lankoides, Wesenbergus, Balkanella and Enter oides. No new generic terms are given by Castellani and Chalmers in their later report (Ann. Inst. Past., 34, 1920, 600). SURVEY OF CLASSIFICATIONS OF BACTERIA 11 Orla-Jensen (Jour. Bact., 6, 1921, 263), in a paper published after the manuscript of the first edition of the Manual was prepared, suggested the use of Colibacterium and Aerogeneshacterium for the two genera in the coliform group and adds quite a number of other new terms formed in accordance with his system of nomenclature. These are, in most cases, synonjTns of earlier valid names. The new terms are Coccomonas, Spiro- monas (used in a new, different sense from that of earlier authors), Fluoro- monas, Photomonas, Propionicoccus, Buty rich sir idium and Putriclostridium. Many new terms are proposed in the classification drawn up by Heller (Jour. Bact., 6, 1921, 521; and 7, 1922, 1). Details are given in the group of anaerobic spore-formers onlj*. Here each of the new generic terms is based on a single species. The following outline is given in the first of these papers, two new genera {Rivoltillus and Metchnikovillus) being made the type genera for two new subfamilies Clostridioideae and Putrificoideae, respectively. Phylum I. Bacteria Class I. Eubacterieae Order 1. Eubacieriales Family C (?). Clostridiaceae Subfamily 1. Clostridioideae Subfamily 2. Putrificoideae Order 2. Thiobacteriales Order 3. Chlamydohacteriales Class II. Myxohacterieae In the more complete outline in the second paper, one generic term (Clostridium) is old, although it is used in a new and restricted sense, while with the exception of the type genera mentioned above, the other terms are new. In the subfamily Clostridioideae, the new terms are Omelianskillus, Macintoshillus, Douglasillus, Henrilliis, Flemingillus, Vallorillus, Multi- fermentans, Hihlerillus, Welchillus, Stoddardillus, Arloingillus, Meyerillus and Novillus. Ten new generic terms are used in the subfamily Putrifi- coideae as follows: Sequinillus, Reglillus, Rohertsonillus, Nicollaierillus, Martellillus, Recordillus, Tissierillus, Putrificus, Ermengemillus, and Wein- bergillus. As there does not seem to be any good reason for sub-dividing the genus Clostridium in this way, the latter term has been used to cover anaerobic spore-forming rods in all previous editions of the Manual, and is again used in the present edition in this sense rather than with the restricted meaning proposed bj^ Heller. Enderlein (Sitzber. Gesell. Naturf . Freunde, Berlin, 1917, 309) proposed an outline classification covering the Kingdom of Mychota, or bacteria, which was based on comparative morphology with special emphasis on life cycles. This was as follows: 12 MANUAL OF DETERMINATIVE BACTERIOLOGY Phylum I. Dimychota Ki reis A. Holocyclomorpha Class I. Gonascota Order a. Synascota Family 1. Schaudinnidae Genus a. Schaudinnum b. Theciobactrum Family 2. Sphaerotilidae Genus a. Phragmidiothrix b. Newskia c. Chlaynydothrix d. Sphaerotilus e. Clonothrix Family 3. Syncrotidae Genus a. Crenolhrix h. Beggiatoa c. Syncroiis d. Zygoslasis Family 4. Spirillidae Genus a. Gallionella b. Spirillum c. Dicrospirillum Family 5. Spirochaeiidae Genus a. Cristispira h. Treponema c. Entomospira d. Spirochaeta e. Cacospira Family 6. Microspiridae Genus a. Spirobacillus b. Spirosoma c. Photobacterium d. Microspira e. Dicrospira Family 7. Corynobacteriidae Subfamily 1. Aclinomycinae Genus a. Actinomyces Subfamily 2. Eisenberginae Genus a. Eisenbergia Subfamily 3. Sclerolrichinae Genus a. Zetlnowia b. Schlerothrix Subfamily 4. Corynobacteriinae Genus a. Corynobaclerium h. Heterocyslia c. Cladascus d. Zygoplagia Subfamily 5. Pseudosirepinae Genus a. Pseudostreplus Order b. Ascoia Family 8. Bacteriidae SURVEY OF CLASSIFICATIONS OF BACTERIA 13 Genus a. Atremis b. Bacterium c. Lamprella d. Eucystia e. Dicrobactrum f. Acysiia Family 9. Fusiformidae Genus a. Fusiformis Class II. Sporascota Order a. Parasynascota Family 10. Migulanidae Genus a. Migulanum Order b. Parascota Family 11. Bacillidae Genus a. Rhagadascia h. Plectridium c. Bacillus d. Bacirillum e. Kochella f. Fischerinum Kreis B. Hemicyclomorpha Class I. Anascola Familj' 12. Hemallosidae Genus a. Hemallosis Phylum II. Monomychota Kreis A. Acyclomorpha Family 1. Mogallidae Genus a. Mogallia Family 2. Sarcinidae Genus a. Diplococcus b. Sarcina c. Paulosarcina Family 3. Micrococcidae Genus a. Micrococcus h. Planococcus c. Streptococcus d. Phacelium Three of the new generic term.s, Cladasciis tj^pe species C. furcabilis Enderlein, Zygoplagia type species Z. alternans Enderlein and Heterocystia type species H. multiformiis Enderlein, had been proposed in an earHer paper (Sitzber. Gesell. Naturf. Freunde, BerUn, 1916, 395). The following generic terms in the 1917 outline are new: Schaudinnum, TJieciobactrum, Syncrotis, Zygostasis, Dicro spirillum, Entomospira, Cacospira, Dicrospira, Eisenhergia, Zettnoivia, Pseudostreptus, Atremis, Lamprella, Eucystia, Dicro- bactrum, Acystia, Migulanum, Rhagadascia, Kochella, Fischerinum, Hemal- losis, Mogallia, Paulosarcina and Phacelium. Note that Corynobacterium is spelled with an o instead of an e. Terms accepted from earlier workers that have not previously been 14 MANUAL OF DETERMINATIVE BACTERIOLOGY mentioned are: Spirobacillus Metschnikoff (Ann. Inst. Past., 3, 1889, 62), Photohacterium Beijerinck, Maanblad voor Natuurwetenschappen Amster- dam, 16, 1889, 1 and Arch. Ne^rl. d. Sci. Exactes, 23, 1889, 401), and Sclerothrix Metschnikoff (Arch. f. Path. Anat. u. Physiol., 113, 1888, 63-94; not Sclerothrix Kuetzing, Species Algarum, 1849, 319). The above outHne was changed in 1925, p. 235 ff. (Bakterien-Cyclogenie, BerHn, 390 pp.) by the addition of one new family, Chondromycidae, to include the genus Newskia, formerly included in Sphaerotilidae, and nine genera not previously given as follows : Chondromyces, Cystodesmia, Mono- cystia, Ophiocystia, Apelmocoena, Polyangium, Cystoecemia, Myxococcus and Dactylocoena. All except Chondromyces, Polyangium and Myxococcus are taken from Enderlein (Bemerkungen zur Systematik der Chondromyciden, Berlin, 1924, 6 pp.). The new genus Lohnisium is added in the Family Eisenhergiinae to in- clude the acetic acid and legume bacteria, and he also proposes the generic term Macrocystita (p. 278) for certain bacteria described by Peklo (0 msici krvave (Study of the blood louse). Zemedelskdho Archivu (Agri- cultural Archives), 1, 1916) from aphids. According to Enderlein it is not clear whether this genus should be included in the Family Bacteriidae or in Corynobacteriidae . Two genera proposed by others are also accepted. These are Calym- matohacterium Aragao and Vianna (Mem. Inst. Oswaldo Cruz, 6, 1912, 211) placed in the family Migulanidae, and Leuconostoc Van Tieghem placed in the family Micrococcidae. Later Enderlein (Sitzber. Gesell. Naturf. Freunde Berlin, 1930, 104-105) accepts Serratia Bizio in place of Dicrohactrum, and Leptotrichia Trevisan in place of Syncrotis. Streptus with Streptus scarlaiinae as type species, is proposed to cover the streptococci not included in Pseudostreptus. The outline suggested by Pringsheim (Lotos, 71, 1923, 357) is similar to that used by Lehmann and Neumann (Atlas und Grundriss der Bakterio- logie, 2 vols., 1896, Miinchen). It is a conventional division into spherical, rod-shaped and curved forms so far as the true bacteria are concerned except that the pseudomonads are included in the same family as the vibrios and spirilla. Rhodohacteriales is recognized as an order to include the sulfur purple bacteria and the nonsulfur purple bacteria. Few details are given in regard to the other orders. His outline follows: Schizomycetes Order I. Eubacteriales Family 1. Coccaceae Genus a. Streptococcus b. Micrococcus c. Sarcina SURVEY OF CLASSIFICATIONS OF BACTERIA 15 Family 2. Bacleriaceae Genus a. Bacterium b. Bacillus Family 3. Spirillaceae Genus a. Pseudomonas b. Vibrio c . Spirillum Order II. Rhodobacteriales Family 1. Rhodobacterinae 2. Thiorhodinae Order III. M yxobacteriales Family 1. Myxobacteriaceae Order IV. Mycobacteriales Family 1. Corynebacteriaceae 2. Mycobacteriaceac 3. Actinomycelaceae (Also possibly the long rod, lactic acid bacteria.) Order V. Desmobacteriales Family 1. C hlamydobacteriaceae 2. Beggiatoaceae The first outline classification drawn up by Janke (Allgemeine Technische Mikrobiologie, I Teil, Dresden, 1924, p. 63) is an adaptation and expansion of that drawn up by Migula (System der Bakterien, 1900) . The new genera recognized by Janke are Planostre^tococcus A. Meyer (Die Zelle der Bak- terien, Jena, 1912), Thioploca Lauterborn (Ber. dtsch. Bot. Gesell,, 25, 1907, 238), Thiohacterium Molisch (Cent. f. Bakt., II Abt., 33, 1912, 55), Thiobacillus Beijerinck (Cent. f. Bakt., II Abt., 11, 1904, 593), Thiovihrio Janke {loc. cit.), Thiospirillum Winogradsky (Beitrage zu Morphol. u. Physiol, d. Bakterien. Heft I. Schwefelbakterien. Leipzig, 1888), Thio- sphaerella Nadson (Bull. Jar. bot. Petersburg, 13, 1913, 106; ref. in Cent. f. Bakt., II Abt., 43, 1915, 469), Thiovulum Hintze (Ber. Dtsch. Bot. Gesell., 31, 1913, 189), Spirophyllum Ellis (Proc. Roy. Soc. Edinburgh, 27, I, 1907, 21; ref. in Cent. f. Bakt., II Abt., 19, 1907, 502), Nodofolium Ellis (Cent. f. Bakt., II Abt., 26, 1910, 321), and Actinococcus Beijerinck (Fol. Microbiol.,:^, 1914, 185). Janke 's outline classification is given below: Order I. Eubacteria Family 1. Coccaceae Genus a. Streptococcus h. Micrococcus c. Sarcina d. Planostreptococcus e. Planococcus f. Planosarcina Family 2. Bacleriaceae Genus a. Bacillus b. Bacterium IQ RIANUAL OF DETERMINATIVE BACTERIOLOGY Family 3. Spirillaceae Genus a. Microspira b. Spirillum c. Spirosoma Order II. Rhodobacteria Family 1. Thiorhodaceae Subfamily la. Thiocysteae Genus a. Thiocystis b. Thiocapsa c. Thiosphaera d. Thiosphaerion e. Thiosarcina Subfamily 2b. Lamprocysteae Genus a. Lamprocystis Subfamily 3c. Thiopedieae Genus a. Thiopedia h. Thioderma Subfamily 4d. Amoebobacterieae Genus a. Amoebobacter b. Thiothece c. Thiodictyon d. Thiopolycoccus Subfamily 5e. Chromatieae Genus a. Chroviatium b. Rhabdochromatium c. Thiorhodospirillu7n Subfamily 6f. Rhodocapseae Genus a. Rhodocapsa b. Rhodothece Family 2. Athiorhodaceae Subfamily la. Rhodocysteae Genus a. Rhodocystis h. Rhodonostoc c. Rhodococcus d. Rhodobacterium e. Rhodobacillus f. Rhodovibrio g. Rhodospirillum Order III. Thiobacteria Family 1. Beggiatoaceae Genus a. Thiothrix h. Beggiatoa c. Thioploca Family 2. Thiobacteriaceae Genus a. Thiophysa h. Thiobacterium c. Thiobacillus d. Thiovibrio e. Thiospirillum f. Thiosphaerella g. Thiovulum h. Achromatium SURVEY OP CLASSIFICATIONS OP BACTERIA 17 Order IV. Phycobacteria Genus a. Leptothrix b. Clonothrix c. Cladothrix d. Crenolhrix e. Phragmidioihrix Appendix Genera Gallionella, Spirophyllum, Nodofolium Order V. Mycobacteria Family 1. Mycobacteriaceae Genus a. Cory neb acterium b. Mycobacterium Family 2. Actinomycetaceae Genus a. Actinomyces b. Actinococcus Order VI. Myxobacteria Familj^ 1. Myxobacteriacene Genus a. Myxococcus b. Chondromyces c. Polyangium Lehmann and Neumann (Bakt. Diag., 2 vols., 7th ed., Miinchen, 1926-27 ; Breed, Eng. trans., New York, 1931) developed their first simple and much used outline classification, dra\vn up in 1896, in later editions of their Determinative Bacteriology. The 1927 Lehmann and Neumann outline is as follows: Class I. Schizomycetes Order I. Schizomycetales Family 1. Coccaceae Genus a. Streptococcus b. Sarcina * c. Micrococcus Sub-genus a. Diplococcus b. (Gram-positive group) Family 2. Bacleriaceae Genus a. Bacterium Sub-genus a. Nitrosomonas b. Nitrobacter c. Rliizobium d. Haemophilus e. Brucella f. Pasteurella g. (Glanders and dysenterj' group) h. (Photogenic group) i. (Aerogenes group) j. Encapsulatus k. (Typhoid group)* 1. Salmonella m. (Coli group)* * In a footnote under these groups, the authors refer to the names given by Castel- lani and Chalmers. 18 MANUAL OF DETERMINATIVE BACTERIOLOGY n. Aceiobacierium o. (Cloacae group) p. (Red chromogens) q. (Blue and violet chromogens) r. Pseudornonas s. Proteus App. Erysipelothrix Genus h. Fusobaclerium c. Plocamobacierium Family 3. Desmobacteriaceae Genus a. Beggiaioa h. Leplothrix Sub i-genus a. Leplothrix b. Chlamydothrix Genus c. Crenothrix d. Cladolhrix e. Thiothrix Family 4. Spirillaceae Genus a. Vibrio b. Spirillum Family 5. Spirochaetaceae Genus a. Spirochaela Family 6. Bacillaceae Genus a. Bacillus Sub i-genus a. (Aerobic group) b. (Anaerobic group) Order II. Actinomyceiales Family 1. Proactinomycelaceae Genus a. Corynebaclerium b. Mycobacterium Family 2. A ctinomycetaceae Genus a. Actinomyces The generic term Bacterium is retained in this outUne to cover those groups of the true bacteria that are Gram-negative, non-spore-forming, motile and non-motile rods. Lehmann and Neumann recognize 20 sub- groups in this genus, many of which correspond with the genera recognized in the Manual. In an effort to develop a rational nomenclature the term Acetobacterium (occurs first in review by Ludwig, Cent. f. Bakt., II Abt., 4, 1898, 870) is used in place of Acetohacter, Plocamohacterium (Loewi, Wien. klin. Wchnschr., 33, 1920, 730) in place of Lactohacillus, and Fusobacterium (Knorr, Cent. f. Bakt., I Abt., Orig., 89, 1922, 4) in place of Fusiformis without regard to priority. Encapsulatus Castellani and Chalmers (Man- ual Tropical Med., 3rd ed., 1919, 934) is used in place of Klebsiella Trevisan (Atti Accad. Fis.-Med.-Stat. Milano, Ser. 4, 3, 1885, 107). Janke (Cent. f. Bakt., II Abt., 66, 1926, 481) reprints the classification developed in the first edition of the present Manual and compares it with that proposed by Orla- Jensen and Enderlein. SURVEY OF CLASSIFICATIONS OF BACTERIA 19 The second complete outline drawn up by Janke (Oesterr. Bot. Zeitschr., 78, 1929, 108) is similar to the classification employed by Lehmann and Xeumann (Bakt. Diag., 2 vols., 7th ed., Miinchen, 1926-27). He follows Enderlein in placing Azotohacter in close association with the spore-forming rods. No new generic terms are suggested. His sub-groups of the genus Bacterium are even more closely similar to the genera used in the present edition of the jNIaxual than are the sub-groups of Lehmann and Neumann. Family 1. Coccaceae Genus a. Micrococcus b. Neisseria c. Streptococcus Divided into 4 groups. d. Sarcina Divided into 2 groups. Family 2. Bacillaceae Genus a. Bacillus Divided into 16 groups, b. Azotohacter Family 3. Bacteriaceae Genus a. Bacterium Divided into 27 groups. b. Fusifortnis Family 4. Corynobacteriaceae Genus a. M ycohacterium h. Corynobacierium c. Actinomyces Family 5. Spirillaceae Genus a. Microspira Divided into 2 groups, b. Spirillum Divided into 2 groups. Family 6. Spirochaetaceae Genus a. Spirochaeta b. Borrelia c. Treponema d. Cristispira e. Saprospira f. Leptospira Family 7. Desmobacleriaceae Genus 1 a. Beggiatoa b. Thioploca c. Thiothrix d. Leptolrichia e. Crenothrix f. Sphaerotilus S- Clonothrix h. i. Leptothrix Phragmidiothrix 20 ' MANUAL OF DETERMINATIVE BACTERIOLOGY Family 8. M yxobacteriaceae Genus a. Myxococcus b. Polyangium c. Chondromyces Pribram (Jour. Bact., 18, 1929, 361) has rearranged some groups and combined others (e.g., Rhizohium, Diplococcus, Leuconostoc, Serratia, Flavohacterium, Chromobacterium, Achromohader, Cellulomonas) recognized in the first edition of the Manual with Httle change in the nomenclature except among the anaerobic non-spore-forming rods and among the spore- forming rods. Unfortunately, he has sometimes used family and species names as generic names, thus in the latter case introducing adjectives and adjectival terms as substantives. New generic terms suggested are: Dialisterea, Bacteroidea, Centrosporus, Fusibacillus, Pseudobacillus, Mega- therium, Flexus, Anthrax, Botulinus, Chauvoea, Botidinea, Putrificus, Welchia, Phleobacterium, Distasoa, Tissieria, and Actinoidomyces. Astasia as it appears in this outline does not appear to be the same as Astasia Meyer (Flora, 84, 1897, 185). Aerobacillus is not synonymous with Aerobacillus Donker (Inaug. Diss., Delft, 1926). Sideromonas is accepted from Cho- lodny (Ber. Deutsch. Bot. Ges., 40, 1922, 326). Pribram's complete outline follows: Class Schizomycetes Subclass A. Prolozoobacteria Order I. Spirochaetales Family 1. Spirochaetaceae Genus a. Spirochaeta b. Treponema c. Spironema Family 2. Crisiispiraceae Genus a. Saprospira h. Cristispira c. Leptospira Subclass B. Eubacleria Order I. Protobacteriales Family 1. Nitrobacteriaceae Related to Pseudomonas Tribe A. H ydrogenomonadae Genus a. H ydrogenomonas b. M ethanomonas c. Carboxyodomonas Tribe B. Niirobactereae Genus a. Niirosomonas h . Nitrobacter Family 2. Thiobacillaceae Tribe A. Thiobacilleae Genus a. Thiobacillus SURVEY OF CLASSIFICATIONS OF BACTERIA 21 Order II. Metahacteriales Family 1. Pseudomonadaceae Tribe A. Spirilleae Genus a. Spirillum Tribe B. Vibrioneae Genus a. Vibrio Tribe C. Pseudomonadeae Genus a. Pseudomonas h. Azoiobacter Connects with Polyangiaceae and Nitrubacteriaceae Family 2. Bacleriaceae Tribe A. Aerobactereae Genus a. Aerobacter h. Escherichia c. Salmonella d. Eberthella e. Proteus Tribe B. Pasteurelleae Genus a. Alcaligenes h. Pasteurella Connects with Pfeifferella c. Hem,ophilus Connects with Dialisler Family 3. Micrococcaceae Tribe A. Streplococceae Genus a. Neisseria • b. Streptococcus Tribe B. Micrococceae Genus a. Micrococcus h. Staphylococcus c. Sarcina Connects with Algubacteria Subclass C. Mycobacteria Order I. Bacteriomycetales Family 1. Leptotrichaceae Tribe A. Acetobactereae Genus a. Acetobacter Connects with Salmonella and Tissieria Tribe B. Leptotricheae Genus a. Kurthia b. Lactobacillus Connects with Corynebacterium c . Leptotrichia Connects with Erysipelothrix Family 2. Bacteroidaceae Tribe A. Dialistereae Genus a. Type species Dialistereavariegata Connects with Distasoa h. Type species Dialisterea variabilis c. Dialisler 22 MANUAL OF DETERMINATIVE BACTERIOLOGY Connects with Hemophilus Tribe B. Bacieroideae Genus a. Type species Bacteroidea multiformis b. Bacter aides Connects with Tissieria c. Type species Bacteroidea fusiformis Order II. Bacilloviycetales Family 1. Bacillaceae Sub-family la. Aerobacilloideae Tribe A. Aerobacilleae Sub-tribe Al. Centrosporineae Genus a. Centrosporus b. Fusibacillus Sub-tribe A2. Aerobacillineae Genus a. Aerobacillus Tribe B. Pseudobacilleae Genus a. Pseudobacillus Sub-family lb. Bacilloideae Tribe A. Bacilleae Sub-tribe Al. Bacillineae Genus a. Bacillus h. Megatherium Sub-tribe A2. Astasineae Genus a. Astasia h. Plexus " Tribe B. Anthraceae Genus a. Anthrax Family 2. Clostridiaceae Sub-family 2a. Botulinoideae Tribe A. Botulineae Genus a. Botulinus b. Chauvoea c. — — Type species Botulinea saccharolytica d. Type species Botulinea butyrica Tribe B. Putrificeae Genus a. Putrificus Sub-family 2b. Clostridioideae Tribe A. Welchieae Genus a. Welchia Tribe B. Clostridieae Genus a. Clostridium , Order III. Actinomycetales Family 1. M ycobacteriaceae Tribe A. Actinobacilleae Genus a. Pfeifferella Connects with Pasteurella b. Actinobacillus c. Corynebacterium d. Erysipelothrix Connects with Leptotrichia SURVEY OF CLASSIFICATIONS OF BACTERIA 23 Tribe B. Mycobactereae Genus a. Phleobacteriuni b. Mycobacterium Tribe C. Tissierieae Genus a. Distasoa b. Tissieria Connects with Bacteroides, Corynebacterium and Acelobacter Family 2. Actinomycetaceae Tribe A. Actinoidomycetaceae Genus a. Actinoidomyces Tribe B. Actinomycetaceae Genus a. Actinomyces Subclass D. Algobacteria Order I. Desmobacteriales Family 1. Sphaerotilaceae Genus a. Sphaerotilus Order II. Siderobacteriales Family 1. Chlamydotrichaceae Tribe A. Chlamydotricheae Genus a. Leptothrix h. Crenothrix Family 2. Siderocapsaceae Genus a. Didymohelix h. Siderocapsa c. Sideromonas Order HI. Thiobacteriales Family 1. Rhodobacteriaceae Sub-family la. Chromatoideae Tribe A. Thiocapseae Genus a. Thiocystis b. Thiosphaera c. Thiosphaerion d. Thiocapsa e. Thiosarcina f. Lamprocysiis Tribe B. Thiopedieae Genus a. Lampropedia h. Thioderma Tribe C. Amoebobacteriae Genus a. Amoebobacter h. Thiodictyon c. Thiothece d. Thiopoly coccus Tribe D. Chromatieae Genus a. Chromatium b. Rhabdomonas c. Thiospirillum d. Rhodocapsa e. Rhodcihece 24 MANUAL OF DETERMINATIVE BACTERIOLOGY Sub-family lb. Rhodobacteroideae Tribe A. Rhodobacteriaceae Genus a. Rhodobacterium b. Rhodobacillus c. Rhodovibrio d. Rhodospirillum 6. Rhodosphaera Tribe B. Rhodocysteae Genus a. Rhodocystis b. Rhodonostoc Connects with Leuconosloc Family 2. Beggiatoaceae Genus a. Thiothrix b. Beggiatoa c. Thioploca Family 3. Achromatiaceae Genus a. Achrcmatium h. Thiophysa c. Thiospira d. Hillhousia Order IV. Myxobacieriales Family 1. Polyangiaceae Genus a. Chondromyces b. Polyangium Family 2. Myxococcaceae Genus a. Myxococcus Later Pribram (Klassification der Schizomyceten (Bakterien), Leipzig and Wien, 1933, 143 pp.) developed this classification into a suggestive out- line based on his experience in caring for the cultures of the Krai Collection. His most interesting contribution is the separation of the class of Schizo- mycetes into three subclasses which are based on differences in fundamental biological and nutritional relationships. The fourth sub-class of his earlier outline (the Protozoohacteria with its single order Spirochaetales) is omitted from this outline. The first class, Algohacteria, includes the bacteria that are primarily free-living in water, usually motile with polar flagellation and live on easily soluble foodstuffs. They are frequently surrounded by in- soluble secretions such as capsules, sheaths, etc., and form insoluble prod- ucts in their protoplasm, such as calcium, sulfur and iron compounds, and pigments. The class Euhacteria includes those bacteria whose normal habitat is the animal body or complex waste products of plant or animal origin. Because of adaptation to environment, these organisms are motile or non-motile and can utilize compounds of complex molecular structure. The third sub-class, Mycobacteria, is adapted to life in soil, and shows a dis- tinct tendency to differentiation in morphology and spore formation. Internationally accepted rules of nomenclature are generally followed, and the generic terms proposed in his earlier outline that were not formed SURVEY OF CLASSIFICATIONS OF BACTERIA 25 in accordance with recommended practices are discarded. He has revived Ulvina Kiitzing, 1837 (status explained by Buchanan, General Systematic Bacteriology, 1925, p. 518) in place of Acetohacter Beijerinck and accepted Plocamohacteriuvi (Loewi, Wien. klin. Wchschr., 38, 1920, 730) in place of Lactobacillus Beijerinck, 1901. Among the spore-forming rods, he has accepted Bactrillum Fischer and Welchillus Heller, 1921. Malleomyces Hallier (Bot. Ztg., 24, 1866, 383) is used for the glanders bacillus. Anthra- cillus is apparentl}' new. The new outline has the following form : Class. Schizomycetes Subclass A. Algobacteria Order 1. Micrococcales Family 1. Micrococcaceae Genus a. Micrococcus b. Rhodococcus c. Rhodocapsa d. Thiocapsa e. Thiosphaera f. Thiosphaerion g- Thiocystis h. Lamprocysiis i. Sarcina J- Thiosarcina Family 2. Fediococcaceae Genus a. Pediococcus b. Lampropedia c. Thiothece d. Thiopolycoccus e. Thioderma f. Amoebomonas to* Rhodothece h. Rhodonostoc i. Thiophysa Order 2. Pseudomonadales Family 1. Pseudomonadaceae Genus a. Pseudomonas b. Rhodobacillus c. Chromatium d. Nitrosomonas e. Vibrio f. Rhodovibrio g- Myxococcus h. Spirillum i. Rhodospirillum J- Thiospira k. Thiospirillum Family 2. Serratiaceae Genus a. Serratia b. Hillhousia 26 MANUAL OF DETERMINATIVE BACTERIOLOGY Family 3. Nitrobacteriaceae Genus a. Nitrobacter b. Rhodobacterium c. Rhodocystis d. Didymohelix e. Sideromonas f. Siderocapsa g. Chondromyces h. Polyangium i. Amoebobacter j. Thiodictyon Family 4. Azotobacteriaceae Genus a. Rhizobium b. Azotobacter Order 3. Leptotrichales Family 1. Leptotrichaceae Genus a. Leptothrix b. Sphaerotilus c. Crenothrix Family 2. Clonothrichaceae Genus a. Clonothrix Order 4. Rhabdonionadales Family 1. Rhabdomonadaceae Genus a. Beggiatoa b. Rhabdomonas c. Thioploca d. Thiothrix Family 2. Spirochaetaceae Genus a. Spirochaeta b. Treponema c. Leptospira d. Cristispira e. Saprospira Subclass B. Eubacteria Order 1. Aerobacteriales Family 1. Aero6ac. of the Schuylkill (River). Synonyms: Pseudomonas capsulala Chester, Man. Dctcrm. Bact., 1901, 322 (Bacillus fluorescens capsulalus Pottien, Ztschr. f. Hyg., 11, 1896, 140); Pseudo- monas dennatogenes Fuhrmann, Cent, f. Bakt., II .\bt., 17, 1906, 356. Short rods, with rounded ends, occur- ring singly, in pairs and in chains. Mo- tile, possessing a polar flagellum. Gram- negative. Gelatin colonies: Grayish-white, trans- lucent, with brownish center, radiate margin, becoming bluish-green. Gelatin stab: Slow crateriform lique- faction, with blue-green fluorescence. Agar slant : Grayish, translucent growth. Medium shows greenish fluores- cent. Broth: Turbid, with slight pellicle and blue-green fluorescence. Stringy sedi- ment. Litmus milk: Coagulated, with slow reduction of litmus; peptonized. Potato: Brownish, spreading, viscid, thick. Indole is formed (trace). Nitrites not produced from nitrates. Aerobic, facultative. Does not grow at 35° to 36°C. Source : Isolated from Schuylkill River water. Habitat: Water. 12. Pseudomonas chlororaphis (Guig- nard and Sauvageau) Bergey et al. (Ba- cillus chlororaphis Guignard and Sauva- geau, Compt. rend. Soc. Biol. Paris, 1, 10 s6r., 1894, 841;. Bergey et al., Manual, 3d ed., 1930, 183; also see Lasseur and Dupaix-Lasseur, Trav. Lab. Microbiol. Fac. Pharm. Nancy, Fasc. 9, 1936, 35.) From Greek, chloros, greenish yellow; r aphis, needle. Rods: 0.8 by 1.5 microns, with rounded ends, occurring singly and in pairs. Mo- tile with polar flagella. Gram-negative. Gelatin colonies: Circular, viscid, transparent, glistening, lobate margin, with fluorescent corona. Dissociates readily (Lasseur and Dupaix-Lasseur, loc. cit.). Gelatin stab: Stratiform liquefaction. Broth: Turbid, fluorescent, with crys- tals of green, water-soluble chloro- raphine. Litmus milk : Coagulation. Peptoniza- tion. Crystals of chlororaphine form in the central part of the culture. Potato: Citron-yellow layer. Crystals of chlororaphine are formed. Nitrates reduced to nitrites. Indole not formed. Pigment formation: Asparagine, potas- sium phosphate, glycerol, sulfate of mag- nesium and sulfate of iron are indispensa- ble to the formation of crystals of chlororaphine. Aerobic, facultative. Optimum tem- perature 25° to 30°C. Pathogenic for laboratory animals. Exotoxin formed. Habitat: Water. 13. Pseudomonas myxogenes Fuhr- mann. (Cent. f. Bakt., II Abt., 17, 1907, 356.) From Greek, myxa, mucus; gennao, to beget; M. L. slime producing. Rods: 0.4 to 0.5 by 1.0 to 1.5 microns, occurring singly and in pairs. Motile, possessing a bundle of five to seven polar flagella. Gram-negative. 94 MANUAL OF DETERMINATIVE BACTERIOLOGY Gelatin colonies: Smooth, soft, flat, spreading, entire, yellowish-green. Gelatin stab: Growth along stab. Liquefaction with yellowish-white sedi- ment. Agar colonies : Circular, raised, smooth, amorphous, entire. Agar slant: Yellowish-white, moist, glistening, becoming light green-fluores- cent. Broth: Turbid, with yellowish-white sediment. Litmus milk: Flocculent precipitation. Slow peptonization with yellow serum. Alkaline. Potato : Dirty yellow to olive, moist, glistening, entire. Indole is formed. Nitrates reduced to nitrites and am- monia. No gas formed. Aerobic, facultative. Optimum temperature 22°C. Source: Isolated from beer. 14. Pseudomonas septica Bergey et al. {Bacillus fluorescens septicus Stutzer and Wsorow, Cent. f. Bakt., II Abt., 71, 1927, 113; Bergey et al.. Manual, 3rd ed., 1930, 169.) From Greek, septikos, putre- factive, septic. Rods: 0.6 to 0.8 by 0.8 to 2.0 microns, occurring singly. Motile with a polar flagellum. Gram-negative. Gelatin stab: Infundibuliform lique- faction. Agar colonies: Circular with opalescent center and transparent periphery. Agar slant : Moderate, undulate margin. Broth: Turbid with fragile pellicle, greenish in upper poution. Litmus milk: Alkaline, coagulated. Blood serum not liquefied. Acid from glucose. Aerobic, facultative. Optimum temperature 20^0. Habitat: Disease of caterpillars. 15. Pseudomonas boreopolis Gray and Thornton. (Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 74.) From Greek, boreas, the North wind; polis, city; M. L. North City. Rods: 0.5 to 1.0 by 2.0 to 3.0 microns, occurring singly and in pairs. Motile with one to five polar flagella. Gram- negative. Gelatin colonies : Liquefied. Gelatin stab : Liquefied. Medium red- dened. Agar colonies: Circular or amoeboid, white to buff, flat to convex, smooth, glistening, translucent border. Agar slant: Filiform, whitish, raised, smooth, glistening, fluorescent. Broth: Turbid. Nitrates reduced to nitrites. Starch not hydrolyzed. Acid produced from glucose. Attacks naphthalene. Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat: Soil. 16. Pseudomonas smaragdina Migula. (Bacillus S7naragdimis foetidus Rei- man, Inaug. Dissertation, Wiirzburg, 1887;Migula,Syst.d. Bakt.,.?, 1900, 890.) From Greek, smaragdinas, green like the smaragdus, the emerald. Small rods, occurring singly. Non- motile. Gram -negative. Gelatin colonies : Small, convex, irregu- lar, whitish with greenish shimmer. Gelatin stab: Slight surface growth. Infundibuliform liquefaction. The liquefied medium becomes light emerald green in color. Agar colonies: Small, brownish-yellow, convex. Agar slant : Abundant growth with greenish fluorescence. Broth: Turbid. Litmus milk: Not coagulated. Potato: Dark brown, becoming choco- late brown. Indole not formed. Nitrates not reduced. The cultures give off an odor resembling jasmine. Aerobic, facultative. FAMILY PSEUDOMONADACEAE 95 Optimum temperature 37°C. Subcutaneous and intravenous inocula- tions into rabbits cause death in 36 to 48 hours. Source : Isolated from nasal secretions in ozena. 17. Pseudomonas chlorina (Frankland and Frankland) Levine and Soppeland. {Bacillus chlorinus G. and P. Frankland, Philos. Trans. Roy. Soc. London, 178, 1887, 274; Bacterium chlorinum Migula, Syst. d. Bakt., 2, 1900, 471; Levine and Soppeland, Bui. No. 77, Iowa State Agri- cultural College, 1926.) From Greek, chloros, greenish yellow. Piods : 0.5 by 1 .5 micron, occurring singly and in short chains. Non-motile. Gram- negative. Gelatin stab : Crateriform liquefaction with green fluorescence. Lemon yellow sediment. Agar colonies : Circular, raised, smooth, amorphous, entire, becoming greenish yellow. Agar slant : Slightly raised, glistening, the medium becoming light greenish yellow. Broth : Moderate turbidity. Dirtj' yellow sediment. No pellicle. Litmus milk: Peptonized. Litmus re- duced. Potato: Scant, olive green growth. Indole formed. Nitrites produced from nitrates. Starch hydrolyzed. Blood serum liquefied in .5 days. Acid from glucose. Aerobic, facultative. Optimum temperature 22°C. Source : Air. 18. Pseudomonas oleovorans Lee and Chandler. (Jour. Bact., 41, 1941, 378.) From M. L. oil destroying. Short rods: 0.5 by 0.8 to 1.5 microns, occurring singly and in pairs. . Motile. Gram-negative. Gelatin stab : No liquefaction after 6 weeks. Gelatin colonies: Up to 1 mm. in diam- eter, fluorescent ; similar to agar colonies. Surface agar colonies : After 24 hours 1 to 2 mm. in diameter, smooth, convex, shin3% opaque, creamy, fluorescent by transmitted light. Edge entire in young colonies. Deep agar colonies: 0.5 by 1.0 to 1.5 mm., lens-shaped, buff-colored, not flu- orescent. Agar slant : Growth raised, smooth, fluorescent, edge erose. Broth : After 24 hours moderate turbid- ity with slight yellowish viscid sediment. No pellicle or ring. Litmus milk: No change. Indole not formed. Potato : Good growth. Nitrites are produced from nitrates. Starch is hydrolyzed. No acid from glucose, lactose, sucrose, galactose, xylose, mannitol, salicin and glycerol. Equally good growth at 25° and 37°C. Aerobic. Distinctive character: The fluorescent quality of the colonies is not imparted to any of the artificial media used. Source : Isolated from cutting com- pound (oil-water emulsion) circulating in a machine shop. The oil in this compound may be utilized as a sole source of energy. Habitat : Probably oil-soaked soils. Abundant in cutting compounds. 19. Pseudomonas incognita Chester. {Bacillus Jiuorescens incognilus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 436; Chester, Determinative Bacteriology, 1901, 323.) From Latin, in, not; cogito, to think; M. L. unknown. Short rods, with rounded ends, occur- ring singly, in pairs and in chains. Mo- tile, possessing a polar flagellum. Gram- negative. Gelatin colonies: Thin, transparent, slightly granular, becoming greenish. Margin undulate. The medium assumes a blue-green fluorescence. 96 MANUAL OF DETERMINATIVE BACTERIOLOGY Gelatin stab : No liquefaction. Agar slant: Thin, moist, translucent, becoming greenish. Broth: Turbid, with pellicle, becoming greenish. Litmus milk: Slightly acid in a month. The litmus is slowly reduced. Potato: Moist, glistening, spreading, brown. Indole is formed (trace). Nitrites are produced from nitrates. Aerobic, facultative. Optimum temperature 35°C. Habitat: Water. 20. Pseudomonas convexa Chester. {Bacillus fluorescens convexus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 438; Chester, Determinative Bacteriology, 1901, 325.) From Latin, convexus, con- vex, arched. Short, thick rods, with rounded ends. Motile, possessing a polar flagellum. Gram -negative. Gelatin colonies: Circular, convex, glistening, bright greenish, translucent. The medium becomes blue-green, flu- orescent. Gelatin stab: Light green, raised, gUstening surface growth. No liquefac- tion. Agar slant : Moist, translucent, glisten- ing, light greenish. The medium as- sumes a greenish color. Broth: Turbid, becoming greenish. Litmus milk: No coagulation; alkaline. Potato: Pale brown, spreading. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30°C. Habitat: Water. 21. Pseudomonas mildenbergii Ber- gey et al. (Der blaue bacillus, Milden- berg. Cent. f. Bakt., II Abt., 56, 1922, 309; Pseudomonas cyanogena Bergey et al.. Manual, 1st ed., 1923, 129; not Bacil- lus cyanogenes Flugge, Die Mikroorgan- ismen, 1886, 201 ; not Pseudomonas cyano- genes Hammer, Dairy Bact., 1928, 70; Bergey et al.. Manual, 3rd ed., 1930, 172.) Named for Mildenberg who first isolated this species. Rods: 0.3 to 0.5 by 1.0 to 3.5 microns, with rounded ends, occurring singly. Motile, possessing polar flagella. Gram- negative . Gelatin colonies: Circular, lobed, smooth, glistening, slightly raised, steel- blue, entire. Gelatin stab: No liquefaction. Agar colonies: Small, circular, yellow- ish or reddish-yellow, entire, becoming lobed, grajash-green, iridescent. The medium becomes dirty grayish-green. Agar slant: Smooth, spreading, slimy, glistening, grayish-green to dark green, fluorescent. Broth: Turbid green, iridescent to opalescent with slimy sediment. Litmus milk: Not coagulated, blue ring. Potato: Slimy, glistening, spreading, steel blue. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25''C. Source: Isolated from air. 2. Pseudomonas putida (Trevisan) Migula. {Bacillus Jluorcscens putidus Flugge, Die Mila-oorganismen, 2 Aufl., 1886, 288; Bacillus putidus Trevisan, I gen. e le specie d. Batteriacee, 1889, 18; Migula, in Engler and Prantl, Die na- tiir. Pflanzenfam., 1, la, 1895, 29; Ba- cillus fluorescens putridus (sic) Kruse, in Flugge, Die Mikroorganismen, 2, 1896, 292; Bacterium putidum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 271 ; Pseudomonas putrida (sic) Migula, Syst. d. Bakt., 2, 1900, 912.) It is not clear which spelling should be used. Either is correct. From Latin putida or putrida, rotten, stinking. Rods, with rounded ends. Motile, FAMILY PSEUDOMONADACEAE 97 possessing polar flagella. Gram-negative. Gelatin colonies : Small, finely granular, fluorescent with dark center, surrounded by a yellow zone, with pale gray margin. Gelatin stab : Dirty-white surface growth, becoming greenish, fluorescent. No liquefaction. Agar colonies : Circular, raised, smooth, amorphous, entire, with fluorescent zone around the periphery. Agar slant : Yellowish-green layer, be- coming fluorescent. Broth: Turbid, fluorescent. Litmus milk: Unchanged. Potato: Thin, gray to brownish, slimj^ layer. Cultures give off odor of trimethyl- amine. Indole not formed. Nitrites are produced from nitrates. Aerobic, facultative. Optimum temperature 25°C. Will grow at 37°C. (Pteid et al., Penn. Agr. Exp. Sta., Bull. 422, 1942, 9). Distinctive features: Identical with Pseudomonas fluorescens Migula accord- ing to Lehmann and Neumann {loc. cit.) except that it does not liquefy gelatin. See Pseudomonas eisenbergii Migula. Habitat; Putrefying materials; water. 23. Pseudomonas scissa (Frankland and Frankland) Migula. {Bacillus scis- 8MS G. and P. Frankland, Ztschr. f. Hyg., 6, 1889, 398; Migula, Syst. d. Bakt., 2, 1900, 927.) From Latin, scissus, p.p. of scindo, to cut. Rods: 0.5 by 0.5 to 1.0 micron, with rounded ends, occurring singly, in pairs and in chains; on gelatin, coccus-like. Motile with presumably polar flagella. Gram-negative. Gelatin colonies: Small, greenish. Gelatin stab : Thin, smooth, ghstening surface growth, irregular, serrate margin. No liquefaction. The medium becomes light green in color. Agar slant: Smooth, glistening, lobed. The medium assumes a greenish color. Broth: Turbid, with whitish sediment. Litmus milk : Not coagulated. Potato : Glistening, reddish-brown growth. Indole not formed. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 20°C. Distinctive characters : Resembles Pseudomonas viscosa Migula. Source : Found in water and soils in Kent, England. Habitat : Water and soil. 24. Pseudomonas ovalis Chester. (Bacillus fluorescens ovalis Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 9; Chester, Determinative Bacteriologj^, 1901, 325; not Bacillus ovalis Wright, Memoirs Nat. Acad. Sci., 7, 1895, 435.) From ovum, egg; M. L. oval. Rods: 0.3 to 0.7 by 0.7 to 1.3 microns, occurring singly. Motile, possessing a single polar flagellum. Gram-negative. Gelatin colonies : Irregular, lobate, slightly granular. Gelatin stab : No liquefaction. Agar colonies : Circular, opaque, entire, ureenish fluorescence. Agar slant: Thick, white, becoming greenish^ fluorescent. Broth: Turbid, with pellicle. Litmus milk : No coagulation ; alkaline. Potato: Luxuriant, dirty-brown. Indole not formed. Nitrites not produced from nitrates. Starch not hydrolyzed. Blood serum not liquefied. Acid from glucose. Aerobic, facultative. Optimum temperature 25°C. Habitat: Soil. Has been found in in- testinal canal. 25. Pseudomonas striata Chester. (Bacillus striatus viridis Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 22; Chester, Determinative Bacteriology, 1901, 325.) From Latin, strio, streak, groove. Slender rods, of variable lengths, stain- MANUAL OF DETERMINATIVE BACTERIOLOGY ing irregularly, occurring singly and in pairs. Motile, possessing polar flagella. Gram -negative. Gelatin colonies: Circular, yellowish, with filamentous border. Gelatin stab: Raised, white surface growth. No liquefaction. Agar slant: Thin, yellowish-green, smooth, glistening. Broth: Turbid, becoming slightly greenish. Litmus milk : No coagulation ; becom- ing alkaline; litmus reduced. Potato : Moist, glistening, becoming chocolate-brown. Indole is formed. Nitrites are produced from nitrates. Aerobic. Optimum temperature 25°C. Habitat: Soil. 26. Pseudomonas denitrificans Ber- gey et al. {Bacillus denitrificans fluo- rescens Christensen, Cent. f. Bakt., II Abt., 11, 1903, 190; Bergey etal.. Manual, 1st ed., 1923, 131.) From Latin, de, from, out of; nitrum, soda, nitre; M. L. denitrifying. Rods: 0.5 to 0.7 by 0.5 to 1.25 microns, occurring singly and in pairs in large, slimy masses. Motile. Gram-negative. Gelatin colonies: Small, circular, con- toured, raised, moist, pearly-gray, glis- tening. Gelatin stab: Whitish, lobed surface growth. Yellowish-green growth in stab. No liquefaction. Agar colonies : Pearly white, circular, entire. Agar slant: Broad, whitish, contoured, moist, entire. Broth: Turbid, with thick, wrinkled pellicle. Litmus milk : Not coagulated. Potato: Reddish-gray layer. Indole not formed. Nitrates reduced with production of nitrogen. Aerobic, facultative. Optimum temperature 25°C. Habitat: Soil. 27. Pseudomonas solaniolens Paine. (Rept. Int. Conf. Phytopath. and Econ. Ent. Holland, 1923, 77; Phytomonas so- laniolens Bergey et al.. Manual, 3rd ed., 1930, 274.) From M. L. Solanum, a generic name. Small oval rods: Motile with a polar flagellum. Gram-negative. Produces an iridescence in gelatin. Gelatin: No liquefaction. Gelatin colonies: Round. Iridescence in medium. Agar colonies : Pale buff. Litmus milk: Curd, with no sign of digestion. Potato : Pale buff-colored growth, no change in medium. Nitrites not produced from nitrates. Acid but not gas from glucose. No acid or gas from lactose, sucrose, manni- tol or glycerol. Starch: Action feeble. Optimum temperature 20° to 30°C. Aerobic, facultative. Source : Isolated from potato showing internal rust spots. 28. Pseudomonas eisenbergii Migula. (Bacillus fluurescens non liquefaciens Eisenberg, Bakt. Diag., 3 Aufl., 1891, 145; Bacillus fluorescens immobilis Kruse, in Fliigge, Die Mikroorganismen, 2, 1896, 294; Migula, Syst. d. Bakt., 2, 1900, 913; Bacterium immobile Chester, Man. De- term. Bact., 1901, 180; Pseudomonas non- liquefaciens Bergey et al., Manual, 1st ed., 1923, 132.) Named for Eisenberg, the bacteriologist who first reported this species. Short, slender rods, with rounded ends, occurring singly. Non-motile. Kruse (loc. cit., p. 293) lists the motile form of this organism as Bacillus fluorescens non liquefaciens. Gram-negative. Gelatin colonies : Fern-like surface colonies. Medium around colonies has a pearly luster. Gelatin stab : Surface growth has fluo- rescent shimmer. No liquefaction. Agar slant : Greenish layer. Broth: Turbid, fluorescent. FAMILY PSEUDOMONADACEAE 99 Litmus milk : Unchanged. Potato : Diffuse, brownish layer. Me- dium acquires a grayish-blue color. Indole is not formed. Nitrites produced from nitrates. Acid from glucose. Blood serum liquefied. Aerobic, facultative. Optimum temperature 25°C. Not pathogenic. Habitat : Water. 29. Pseudomonas putrefaciens (Derbj^ and Hammer) Long and Hammer. (Achromobacter putrefaciens Derby and Hammer, Iowa Agr. Exp. Sta. Res. Bui. 145, 1931, 401; Long and Hammer, Jour. Bact., 41, 1941, 100.) From Latin, piiirida, rotten, stinking; faciens, mak- ing. Rods: 0.5 to 1.0 by 1.1 to 4.0 microns, occurring singly and in pairs. Motile, with a single flagellum. Gram-negative. Gelatin stab : Rapid, saccate to strati- form liquefaction, with reddish-brown sediment in the liquefied portion. Agar colony : Circular, smooth, glisten- ing, slightly raised, somewhat trans- parent, with brownish tinge. Agar slant : Echinulate, slightly red- dish-brown, viscous. Broth : Turbid, with thin, gray pellicle, and reddish-brown sediment. Litmus milk : Rapid reduction and proteolysis with odor of putrefaction. Potato : Echinulate, smooth, glistening, viscous, reddish-brown. Indole not produced. Nitrites are produced from nitrates. Acid from maltose and sucrose. No action on glucose, fructose, galactose, arabinose, lactose, raffinose, dextrin, inulin, salicin, amygdalin, glycerol. Ammonia is formed. Aerobic, facultative. Optimum temperature 21°C. Source : Isolated from tainted butter. Habitat : IVIilk, cream, butter, water, soil, creamery equipment (Long and Hammer, loc. cit. ; Claydon and Hammer, Iowa Agr. Exp. Sta., Res. Bull. 267, 1939.) 30. Pseudomonas mephitica Claydon and Hammer. (Jour. Bact., 37, 1939, 254.) From Latin, mephitis, pestilential, malo- dorous; M.L. Mephitis, a generic name. Rods : 0.5 to 1.0 by 1.5 to 14.0 microns, occurring singly, in pairs and in chains. Actively motile with a polar flagellum. Gram-negative. Gelatin: Slow liquefaction. Agar colonies: Convex, circular, about 3 mm. in diameter, shiny, grayish white, edge entire, of the consistency of bread dough. Agar slant : Growth grayish-white, wrinkled, echinulate. After 1 or 2 days, a skunk-like odor develops. Broth: Turbid. Sediment. White pellicle. Potato: Growth echinulate, shiny, brownish. Litmus milk: A skunk-like odor devel- ops in 1 to 2 days. Grayish blue surface ring in about 3 days. Alkaline in 7 to 10 daj's. In two weeks complete reduction. Slight proteoh^sis and viscosity. Hydrogen sulfide not produced. Indole not formed. Nitrites produced from nitrates. Acid but not gas produced slowly from glucose, fructose, maltose, and sucrose. No acid from arabinose, dextrin, galac- tose, glycerol, lactose, mannitol, raffinose or salicin. Aerobic, facultative. Optimum temperature 21°C. Growth slight at 5° and 30°C. No growth at 37°C. Source : Several cultures isolated from butter having a skunk-like odor. Habitat : Probably from water. 31. Pseudomonas geniculata (Wright) Chester. {Bacillus geniculatus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 459; Chester, Man. Determ. Bact., 1901, 313; Achromobacter geniculatum Bergey et al.. Manual, 1st ed., 1923, 146.) From Latin, 100 MANUAL OF DETERMINATIVE BACTERIOLOGY geniculatus, p.p. of geniculo, knotted, jointed. Medium-sized rods, occurring singly, in pairs and chains, motile, possessing polar flagella. Gram-negative. Gelatin colonies: Circular, whitish, translucent. Deep colonies yellowish. Gelatin stab : Infundibuliform liquefac- tion. Sediment light pink. Agar slant : Grayish, glistening, trans- lucent, limited, becoming brownish-graj'. Broth : Turbid, with slight gray pellicle and sediment. Litmus milk : Alkaline ; reduction of litmus; slight coagulation. Potato: Thin, brownish, moist, glis- tening, viscid. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat : Water. 32. Pseudomonas fragi (Eichholz) Huss emend. Hussong, Long and Hammer. (Bacterium fragi Eichholz, Cent. f. Bakt., II Abt., 9, 1902, 425; Huss, Cent, f. Bakt., II Abt., 19, 1907, 661; Hussong, Long and Hammer, Iowa Agr. Exp. Sta. Res. Bull. 225, 1937, 122; also see Long and Hammer, Jour. Dairy Sci., 20, 1937, 448.) From Latin /ra^um,. strawberry. Description from Hussong, Long and Hammer, loc. cit. Rods: 0.5 to 1.0 by 0.75 to 4.0 microns, occurring singly, in pairs and in chains. Motile with a polar flagellura. Gram- negative. Gelatin: Crateriform to stratiform liquefaction in 3 to 4 days. Agar colonies: Convex, glistening, gen- erally butyrous, occasionally viscid. Rough, smooth and intermediate forms are recognized in the description quoted. The rough forms are less proteolytic, and less active in the hydrolysis of fats. Agar slant : Growth abundant, spread- ing, raised, white, shiny, generally butyrous. Sweet ester-like odor resem- bling that of the flower of the May apple. Broth : Turbidity and sediment with a thin pellicle. Litmus milk : Acid ring followed by acid coagulum at surface. Complete coagulation in 2 to 3 weeks, some diges- tion. Characteristic May apple or straw- berry odor. Potato : Growth echiuulate to arbores- cent, raised, glistening, white, becoming brownish. Indole not produced. Nitrites not produced from nitrates. Ammonia produced from peptone. Hydrogen sulfide not produced. Acid from glucose and galactose, sometimes arabinose. No acid from glycerol, inulin, lactose, fructose, mal- tose, mannitol, raflfinose, salicin and sucrose. No acetylmethylcarbinol produced. Fat is generally hydrolyzed. Aerobic. Grows from 10° to 30°C. No growth at 37°C. Very sensitive to heat. Source: Isolated from milk and other dairy products, dairy utensils, water, etc. Habitat : Soil and water. Widely dis- tributed (Morrison and Hammer, Jour. Dairy Sci., U, 1941, 9). Hussong (Thesis, Iowa State College, 1932) regards Bacterium fragi Eichholz (loc. cit.) as the R type, Pseudomonas fragariae I Gruber (Cent. f. Bakt., II .\bt., 9, 1902, 705) as the O form, and Pseudomonas fragariae II Gruber (Cent, f. Bakt., II Abt., 14, 1905, 122) as the S form of the same organism. He makes no mention of Pseudomonas fragaroidea Huss (loc. cit.) which from its description would belong to the smooth type. A brief characterization of each of these organisms follows : (1 ) Bacterium fragi came from milk as drawn from an indi- vidual cow; it does not liquefy gelatin, exhibits no fluorescence, is strongly alka- line in litmus milk, and does not grow at 37°C, (2) Pseudomonas fragariae I came from fodder beets; it does not liquefy gelatin, has weak blue-greenish fluorescence, is weakly alkaline in milk, and grows at 37°C, (3) Pseudomonas fragariae II came from pasteurized milk ; it liquefies gelatin, coagulates milk, and does not grow at 37''C, (4) Pseudomonas FAMILY P8EUDOMONADACEAE 101 fragaroidea came from butter; it liquefies gelatin, coagulates milk, and grows at •33. Pseudomonas nebulosa (Wright) Chester. {Bacillus nebulosus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 465; Chester, Man. Determ. Bact., 1901, 311; Achromobacter nehulosum Bergey et al., Manual, 1st ed., 1923, 145; not Bacillus nebulosus Halle, These de Paris, 1898; not Bacillus nebulosus Vincent, Ann. Inst. Past., 21, 1907, 69; not Bacillus nebulosus Migula, Syst. d. Bakt., 2, 1900, 844; not Bacillus nebulosus Gores- line, Jour. Bact., 27, 1934, 52.) From Latin, nebula, mist. Medium-sized rods, occurring singly. Motile, possessing polar flagella. Gram- negative. Gelatin colonies: Thin, circular, gray, translucent, hazy, with white center. Gelatin stab: Crateriform liquefaction. Agar slant: Thin, transparent streak. Broth: Turbid, with graj' sediment. Litmus milk: Alkaline; reduction of litmus. Potato: Scanty growth. Indole not formed. Nitrites not produced from nitrates. Sugar gelatin in deep stab : Fair growth, with some gas formation. Aerobic, facultative. Optimum temperature 30° to 35°C. Habitat : Water. Probable synonym : Pseudomonas cen- Irifugans Chester. (Man. Determ. Bact., 1901, 312; Bacillus centrifugans Wright, Mem. Nat. Acad. Sci., 7, 1895, 462.) 34. Pseudomonas coadunata (Wright) Chester. (Bacillus coadunatus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 460; Chester, iVIan. Determ. Bact., 1901, 310; Achromobacter coadunatum, Bergey et al., ^Lanual, 1923, 147.) 1st ed.. From Latin, coadunatus, to unite closely. Medium-sized rods, with rounded ends, occurring singly, in pairs and in chains. Motile, possessing a polar flagel- lum. Gram -negative. Gelatin colonies: Circular, brownish, dense. Gelatin stab : Crateriform to stratiform liquefaction. Agar slant : Gray, translucent, spreading. Broth: Turbid, with gray pellicle and sediment. The medium has a slight greenish tint. Litmus milk: Acid; coagulated. Indole is formed. Nitrites not produced from nitrates. Sugar gelatin in deep stab : Good growth of discreet and confluent whitish colonies. Marked gas production; no liquefaction. Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat : Water. 35. Pseudomonas multistriata (Wright) Chester. {Bacillus multistriaius\N right. Memoirs Nat. Acad. Sci., 7, 1895, 462; Chester, Man. Determ. Bact., 1901, 310; Achromobacter multistrialum Bergey et al.. Manual, 1st ed., 1923, 147.) From Latin, multus, many, much; striatus, grooved. Medium-sized rods, with rounded ends, occurring singly and in pairs. Motile, possessing polar flagella. Gram-negative. Gelatin colonies : Circular, grayish- white, translucent. Gelatin stab : Crateriform liquefaction. Agar slant : Narrow, translucent, gray- ish streak. Broth: Turbid. Litmus milk : Slightly acid; coagulated. Potato : Grayish to creamy, thick, glis- tening, viscid, spreading. Indole not formed. Nitrites not produced from nitrates. Sugar gelatin in deep stab : Vigorous growth with marked gas production ; also liquefaction. * Prof. E. R. Hitchner, Univ. of Maine, Orono, Maine assisted in rearranging the descriptions of the acid and gas producing pseudomouads {Aeromonas) , April, 1943. 102 MANUAL OF DETERMINATIVE BACTERIOLOGY Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat : Water. 36. Pseudomonas punctata (Zimmer- mann) Chester. (Bacillus punctatus Zimmermann, Bakt. unserer Trink- unci Nutzwasser, Chemnitz, 1, 1890, 38; Bacillus aquatilis communis Kruse, in Fliigge, Die Mikroorganismen, 2, 1896, 315; Bacterium -punctatum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 238; Chester, Man. Determ. Bact., 1901, 313; Achromobacter punctatum Bergey et al., ]VIanual, 1st ed., 1923, 147.) From Latin, punctus, a puncture, point; M. L. punctate, dotted. Rods: 0.7 by 1.0 to 1.5 micron, occurring singly, in pairs and in chains. Motile with a single polar flagellum. Gram- negative. Gelatin colonies: Small, circular, gray, erose to filamentous, punctiform. Gelatin stab: Crateriform liquefaction. No pellicle. Agar slant: Gray, smooth, filamentous. Broth: Turbid with delicate pellicle. Litmus milk: Acid; coagulated; pep- tonized. Potato : Brownish-yellow to brownish- red color. Indole is formed. Nitrites not produced from nitrates. Hydrogen sulfide is formed. Acid and gas from glucose. Aerobic, facultative. Optimum temperature 25° to 30°C. Source : Common in the Chemnitz tap water. Habitat : Cause of a hemorrhagic septi- cemia in carp {Cyprinus) (Schaperclaus, Ztschr. f. Fischerei, 28, 1930; Cent. f. Bakt., II Abt., 105, 1942, 49). 37. Pseudomonas hydrophila (Ches- ter) comb. nov. {Bacillus hydrophilus fuscus Sanarelli, Cent. f. Bakt., 9, 1891, 222; Bacterium hydrophilus fuscus Ches- ter, Delaware College Agr. Expt. Sta., 9th Ann. Rept., 1897, 92; Bacillus hydro- philus Chester, Manual Determ. Bact., 1901, 235; Bacterium hydrophilum Weldin and Levine, Bact. Abs., 7, 1923, 14; Proteus hydrophilus Bergey et al.. Man- ual, 1st ed., 1923, 211; Aeromonas hydro- phila Stanier, Jour. Bact., 46, 1943, 213.) From Greek, hydor, water, philus, loving; M. L. water-loving. It was reported by Russell, Jour. Amer. Med. Assoc, 30, 1898, 1442 and later by Emerson and Norris, Jour. Exper. Med., 7, 1905, 32 who made a complete study of its properties and its pathogenic action. Weldin (Iowa State College Jour. Sci., 1, 1927, 151) considers Bacillus ranicida Ernst (Beitrage z. path. Anat. u. z. Allgemein. Pathol., 8, 1890, 204; Bac- terium ranicida Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 141) a possible synonym of Proteus hydrophilus. Rods: 0.6 by 1.3 microns, occurring singly and in chains. Motile, with a single polar flagellum (Kulp and Borden, Jour, of Bact., U, 1942, 673). Gram- negative. Gelatin colonies: Small, circular, gray, translucent, stippled. Gelatin stab: Napiform liquefaction. Agar colonies : Whitish, raised, moist, stippled. Agar slant : Thin, whitish, glassy, spreading, becoming yellowish. Broth : Turbid, with heavy pellicle. Litmus milk: Acid; coagulated; pep- tonized. Potato: Yellowish-brown, moist slightly raised. Indole is formed. Nitrites produced from nitrates. Acid and gas from glucose, maltose, sucrose and mannitol. No action on lac- tose. Gas ratio H.rCO. = 1:4.71. Methyl red negative, acetylmethylcarbinol posi- tive, indol negative, citrate positive (Speck and Stark, Jour. Bact., U, 1942, 697). Aerobic, facultative. Optimum temperature 37°C. Pathogenic for frogs, salamanders, fish, t'AMILY PSEtTDOMONADACEAE 103 mice, guinea pigs and rabbits, causing hemorrhagic septicemia. Distinctive characters: Much like Pseudo/nonas punctata (Guthrie and Hitchner, Jour. Bact., 43, 1943, 52). Source : Isolated from frogs dead of septicemia (red leg). Habitat : Water and infected fresh water animals. 38. Pseudomonas ichthyosmia (Ham- mer) comb. nov. (Bacillus ichthyosmius Hammer, Iowa Agr. Sta. Res. Bui. 38, 1917; Escherichia ichthyosmia Bergey et al., Manual, 1st ed., 1923, 201; Proteus ichthyosmius Bergey et al.. Manual, 4th ed., 1934, 364.) From Greek, ichthys, a fish; osme, an odor. Rods: 0.6 to 0.8 by 1.0 to M microns, occur singh'. Motile with a single polar flageilum (Breed). Gram-negative. Gelatin stab: Liquefaction. Agar colonies: Small, white, becoming darker with age. Agar slant : Dirty white, viscid growth. Broth : Turbid with gray sediment. Litmus milk: Acid. Litmus reduced. Cultures have fishy odor. Potato : Thin, glistening layer. Indole is formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, maltose, sucrose, glycerol, salicin and mannitol. Lactose, dulcitol, rafhnose and inulin not fermented. Aerobic, facultative. Optimum temperature 20^C. Source : Isolated from can of evaporated milk having a fishy odor. Habitat: Not known. 39. Pseudomonas ambigua (Wright) Chester. (Bacillus ambiguus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 439; Chester, Man. Determ. Bact., 1901, 308; Achromobacler ambiguum Bergey et al.. Manual, 1st ed., 1923, 148.) From Latin, ambiguus, doubtful, uncertain. Small rods, with rounded ends, occur- ring singly, in pairs and in chains. Mo- tile, possessing a polar flageilum. Gram- negative. Gelatin colonies : Gray, translucent, slighth' raised, irregular, radiate, with transparent margin. Gelatin stab: No liquefaction. Agar slant : Gray, limited, entire. Broth: Turbid, with gray sediment. Litmus milk: Acid, slowly coagulated. Potato : Gray to creamy, viscid, spreading. Indole is formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30° to 35°C. Habitat: Water. 40. Pseudomonas sinuosa (Wright) Chester. (Bacillus sinuosus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 440; Chester, Man. Determ. Bact., 1901, 307; Achromobacler sinosum (sic) Bergey et al.. Manual, 1st ed., 1923, 148.) From Latin, sinuosus, full of bends, sinuous. Medium-sized rods, with rounded ends, occurring singly, in pairs and in chains, ^lotile, possessing two to four polar flagella. Gram-negative. Gelatin colonies : Thin, translucent, irregular, center brownish. Gelatin stab: Grayish-white, glisten- ing, translucent. No liquefaction. Agar slant : Scanty, grayish growth. Broth : Turbid, with gray sediment. Litmus milk: Unchanged. Potato : Grayish-white, moist, spread- ing. Indole is formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30° to 35°C. Habitat : Water. 41. Pseudomonas cruciviae Gray^ and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 91; Achro- mobacter cruciviae Bergey et al.. Manual, 3rd ed., 1930, 218.) From Latin, crux, a cross, via, way, road; from Waycross, a place name. 104 MANUAL OF DETERMINATIVE BACTERIOLOGY Rods: 1.0 by 1.0 to 3.0 microns, occur- ring singly and in pairs. Motile with one to five polar flagella. Gram-negative. Gelatin colonies: Circular, white with buff center, convex, smooth, undulate. Gelatin stab : No liquefaction. Agar colonies : Circular or amoeboid, white to buff, flat to convex, smooth, entire. Agar slant: Filiform, pale buff, raised, smooth, undulate. Broth : Turbid. Nitrites not produced from nitrates. Starch not hydrolyzed. No acid in carbohydrate media. Attack phenol and m-cresol. Aerobic, facultative. Optimum temperature 30 to 35°C. Habitat : Soil. 42. Pseudomonas rugosa (Wright) Chester. {Bacillus rugosus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 438; not Bacillus rugosus Henrici, Arb. Bakt. Inst. Tech. Hochsch. Karlsruhe, 1, 1894, 28; not Bacillus rugosus Chester, Deter- minative Bacteriology, 1901, 220; Ches- ter, Determinative Bacteriology, 1901, 323.) From Latin, rugosus, wrinkled. Small rods, with rounded ends, occur- ring singly, in pairs and in chains. Mo- tile, possessing one to four polar flagella. Gram-negative. Gelatin colonies : Grayish, translucent, slightly raised, irregular, sinuous, radi- ately erose to entire. Gelatin stab: Dense, grayish-green, limited, wrinkled, reticulate surface growth. No liquefaction. Agar slant: Grayish-white, limited, slightly wrinkled, translucent. Broth: Turbid, with grajdsh pelUcle and sediment. Litmus milk: Acid, coagulated. Potato: Moist, glistening, brown. Indole is formed. Nitrites not produced from nitrates. Aerobic. Optimum temperature 30°C. Habitat: Water. 43. Pseudomonas desmolyticum Gray and Thornton. (Gray and Thornton, Cent. f. Bakt., II Abt., 7S, 1928, 90; Achromobacter desmolyticum Bergey et al.. Manual, 3rd ed., 1930, 217.) From Greek desinos, bond, band; lytikos, able to dissolve. Rods : 0.7 to 0.8 by 2.0 to 3.0 microns, occurring singly and in pairs. Motile, with one to five polar flagella. Gram- negative. Gelatin colonies: Circular, gray to buff, raised or umbonate. Smooth, glis- tening, entire. Gelatin stab: No liquefaction. Agar colonies: Circular or amoeboid, whitish, flat or convex, smooth, translu- cent to opaque, entire. Agar slant: Filiform, pale buff, raised, smooth, undulate. Broth : Turbid. Nitrites produced from nitrates. Starch not hydrolyzed. Acid from glucose. Attack phenol and naphthalene. Aerobic, facultative. Optimum temperature 25°C. Habitat: Soil. 44. Pseudomonas rathonis Gray and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 90; Achromo- bacter rathonis Bergey et al., Manual, 3rd ed., 1930, 216.) From M. L. of Ratho Park (Edinburgh). Small rods: 0.5 to 1.0 by 1.0 to 3.0 microns. Motile, with polar flagella. Gram-negative. Gelatin colonies: Circular, white, raised, smooth, glistening, undulate. Gelatin stab: No Uquefaction. Agar colonies: Circular, buff, flat, smooth, glistening, entire. Agar slant : Filiform, pale buff, convex, smooth, glistening, undulate. Broth : Turbid, with peUicle. Nitrites produced from nitrates. Starch hydrolyzed. Acid from glucose and glycerol. FAMILY PSEUDOMONADACEAE 105 Attack phenol and cresol at times, also naphthalene. Aerobic, facultative. Optimum temperature 25°C. Habitat: Manure and soil. 45. Pseudomonas dacunhae Gray and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 90; Achro- mobacter dacv7ihae Bergey et al., Manual, 3rd ed., 1930, 217.) From M. L. from the Island of d'Acunha. Rods: 0.5 to 0.8 by 1.5 to 3.0 microns. Motile with one to six polar flagella. Gram-negative. Gelatin colonies : Circular, whitish, raised, smooth, glistening, undulate. Gelatin stab: Xo liquefaction. Agar colonies: Circular to amoeboid, white, flat, glistening, opaque, entire. Agar slant: Filiform, pale buff, raised, smooth, glistening, undulate. Broth: Turbid. Nitrites produced from nitrates. Starch not hydrolyzed. No acid from carbohydrate media. Attack phenol. Aerobic, facultative. Optimum temperature 25°C. Habitat : Soil . 46. Pseudomonas arvilla Gray and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 90; Achromo- bacter arvillum Bergey et al., Manual, 3rd ed., 1930, 217.) From Latin, arvum, an arable field; M. L. dim. a little field. Rods: 0.5 to 0.7 by 2.0 to 3.0 microns. Motile with one to five polar flagella. Gram-negative. Gelatin colonies: Circular, whitish, convex, smooth, glistening, lobate. Gelatin stab: No liquefaction. Agar colonies: Circular or amoeboid, white to buff, flat to convex, smooth, glistening, opaque, entire. Agar slant : Filiform, whitish, concave, smooth, ringed, entire. Broth : Turbid. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid from glucose. Attacks naphthalene. Aerobic, facultative. Optimum temperature 25°C. Habitat: Soil. 47. Pseudomonas salopium Gray and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 91; Achromo- bacter salopium Bergey et al.. Manual, 3rd ed., 1930, 219.) From Latin, Salop, Shropshire. Rods: 0.7 to 1.0 by 1.0 to 3.0 microns, occurring singly and in pairs. Motile with one to six polar flagella. Gram- negative. Gelatin colonies: Circular, grayish- buff, flat, rugose or ringed, translucent border. Gelatin stab: No liquefaction. Agar colonies : Circular or amoeboid, white to buff, flat to convex, smooth, glistening, translucent border, entire. Agar slant: Filiform, whitish, raised, smooth, glistening, lobate. Broth: Turbid with pellicle. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid from glucose and sucrose. Attacks naphthalene. Aerobic, facultative. Optimum temperature 25°C. Habitat: Soil. 48. Pseudomonas minuscula McBeth. (McBeth, Soil Science, 1, 1916, 437; Cellidomonas minuscula Bergey et al.. Manual, 1st ed., 1923, 162.) From Latin dim. rather small. Rods: 0.5 by 0.9 micron. Motile with one to two polar flagella. Gram-negative. Gelatin stab : Moderate growth . Slight napiform liquefaction. Agar colonies: Small, circular, slightly convex, butyrous becoming brittle, gray- ish-white, finely granular, entire. Agar slant: Moderate, flat, grayish- white. Broth : Turbid. 106 MANUAL OF DETERMINATIVE BACTERIOLOGY Litmus milk : Acid, not digested. Potato: No apparent growth. Indole is formed. Nitrites produced from nitrates. Ammonia is produced. Acid from glucose, lactose, maltose, sucrose and starch. Aerobic, facultative. Optimum temperature 20°C. Habitat : Soil. 49. Pseudomonas tralucida Kellerman et al. (Kellerman, McBeth Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 37; Cellulomonas tralucida Bergey et al., Manual 1st ed., 1923, 163.) From Latin, clear, transparent. Rods : 0.6 by 1.2 microns. Motile with one or two polar flagella. Gram-negative. Gelatin stab : No liquefaction. Agar slant: Scant, grayish growth. Broth : Turbid. Litmus milk : Acid. Potato: No growth. Indole not formed. Nitrites produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Habitat: Soil. 50. Pseudomonas mira McBeth. (Mc- Beth, Soil Science, 1, 1916, 437; Cellu- lonwnas mira Bergey et al.. Manual, 1st ed., 1923, 165.) From Latin, minis, won- derful, extraordinary. Rods : 0.4 by 1.6 microns. Motile with a single polar flagellum. Gram-negative. Gelatin stab: Good growth. No lique- faction. Agar colonies : Circular, convex, gray- ish-white, granular, lacerate. Agar slant: Moderate, flat, grayish- white, somewhat iridescent. Broth: Turbid. Litmus milk: Alkaline. Potato: Moderate, grayish-white. Indole not formed. Nitrites produced from nitrates. Ammonia is produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Habitat: Soil. 51. Pseudomonas lindneri Kluyver and Hoppenbrouwers. (Lindner, 50 Ju- bilaumsber. Westpreuss. Bot.-Zool. Vereins, 1928, 253; T ermohacterium mo- bile Lindner, Atlas d. Mikrosk. Grundl. d. Garungsk., 3 Aufl., 2, 1928, Taf. 68; Kluyver and Hoppenbrouwers, Arch. f. Mikrobiol., 2, 1931, 259; Achromobacter mobile Kluyver and Hoppenbrouwers, ibid., 258; not Pseudomonas mobilis Migula, Syst. d. Bakt., 2, 1900, 923.) Named for Lindner, the German bacteri- ologist who first studied this organism. Short rods 1.4 to 2.0 by 4.0 to 5.0 mi- crons. Occurring singly, in pairs and short chains. Motile with a single polar flagellum. Gram-negative. Peptone gelatin: Poor growth. Peptone agar: Poor growth. Wort agar: White, round, raised colo- nies, 1 mm. in diameter. Good growth. Still better where 2 per cent sucrose, or yeast extract with sucrose is added. Chalk added to neutralize acid. Broth : Poor growth in peptone or yeast extract broth unless sugars are added. Carbon dioxide, ethyl alcohol and some lactic acid produced from glucose and fructose, but not from mannose. May or may not ferment sucrose. May pro- duce as much as 10 per cent alcohol. Catalase produced. Anaerobic, facultative. Optimum temperature 30°C. Distinctive character: The fermenta- tion resembles the alcoholic fermentation produced by yeasts. Source : Isolated from the fermenting sap (pulque) of Agave americana in Mexico. Habitat: Fermenting plant juices in tropical countries (Mexico). FAMILY PSEUDOMOXADACEAE 107 52. Pseudomonas membranoformis (Zobell and Allen) Zobell. (Achromo- bacter membranoformis Zobell and Allen, Jour. Bact., 29, 1935, 246; Zobell, Jour. Bact., 46, 1943, 45). From Latin jnem- brana, membrane, and forma having the form of. Rods: 0.9 to 1.2 by 3.5 to 4.8 microns, occurring singly and in pairs. Motile with lophotrichous flagella. Encapsu- lated. Gram-negative. Gelatin stab: Growth fihform, best at top, with slow crateriform licjuefaction. Agar colonies: Circular, 1.0 to 2.5 mm, with crinkled surface. Agar slant : Moderate, beaded, raised growth. Membranous consistency. Be- comes browned with age. Broth : Slight turbidity, tl(jcculent sedi- ment, film of growth on walls of test tube. iNIilk: Xo growth. Potato: Xo growth. Indole not formed. Nitrites not produced from nitrates. No H2S produced. Acid but not gas from glucose, sucrose, dextrin and mannitol. Xo acid from lactose or xylose. Xo diastatic action. Optimum temperature 20° to 25°C. Aerobic. Source : Sea water. Habitat : Sea water. 53. Pseudomonas marinoglutinosa (Zobell and Allen) Zobell. (Achromo- bacter marinoglutinosus Zobell and Allen, Jour. Bact., 29, 1935, 246; Zobell, Jour. Bact., 46, 1943, 45). From Latin marinas, pertaining to the sea, and glutinosus, full of glue, sticky. Short rods: 0.7 to 1.0 by 1.8 to 2.4 mi- crons, with rounded ends, occurring singly, in pairs and in clumps. Motile with polar flagella. Staining granular. Encapsulated. Gram-negative. Gelatin stab : Moderate filiform growth with slight napiform liquefaction. Xo pigment. Agar colonies : Ptound with concentric circles and crinkled radial lines, 1.5 to 5.0 mm. in diameter. No pigment. Agar slant: Moderate, filiform, flat. Butyrous consistency. Broth: Moderate clouding, marked ring, adherent film of growth on test tube wall, and flaky sediment. Milk : No growth. Potato: Xo growth. Indole not formed. Xitrites not produced from nitrates. Hydrogen sulfide and ammonia pro- duced from Bacto-tryptone. Acid but not gas from xylose and dex- trin. Xo acid from glucose, lactose, sucrose and mannitol. Starch is hydrolyzed. Optimum temperature 20° to 25°C. Aerobic, facultative. Source : Sea water. Habitat: Sea water. 54. Pseudomonas gelatica (Gran) Bergey et al. (Bacillus gelaticus Gran, Bergens Museums Aarbog., 1902, 14; Bacterium gelaticum Lundestad, Cent, f. Bakt., II Abt., 75, 1928, 328; Bergey et al., Manual, 3rd ed., 1930, 175.) From French, like gelatin. Pods, ^vith rounded ends, 0.6 to 1.2 by 1.2 to 2.6 microns, occurring singly, in pairs, and sometimes in short chains. Motile. Gram-negative. Fish-gelatin colonies : Circular, trans- parent, glistening, becoming brownish in color. Fish-gelatin stab : Liquefaction infun- dibuliform, with greenish color. Sea-weed agar colonies : Circular, flat, entire, glistening, reddish-brown center with gra.vish -white periphery. Lique- fied. Fish-agar slant: Flat, transparent streak, with undulate margin, reddish- brown . Broth : Turbid with flocculent pellicle, and greenish-yellow sediment. Indole not formed. Xitrites are produced from nitrates. Starch hydrolyzed. 108 MANUAL OF DETERMINATIVE BACTERIOLOGY No action on sugars. Anaerobic, facultative. Optimum temperature 20 to 25°C. Habitat : Sea water of Norwegian coast. 55. Pseudomonas calcis (Drew) Kellerman and Smith. {Bacterium cal- cis Drew, Yearbook Carnegie Inst. Wash., 11, 1912, 136-144; Kellerman and Smith, Proc. Nat. Acad. Sci., 4, 1914, 400.) From Latin calx (calc-), lime. Ovoid rods, 1.1 by 1.5 to 3 microns, usually single but may form long chains. Actively motile with one polar flagellum. Gram-negative. Grows best in sea water or 3 per cent salt media. Deposits CaCOs. Agar colonies : Circular, with finely irregular outline, granular appearance, elevated, spreading; old colonies having brownish tinge in center. Gelatin stab : Infundibuliform lique- faction. Gelatin colonies: Small, with lique- faction. Broth : Good growth especially in pres- ence of potassium nitrate, peptone or calcium malate. Acid from glucose, mannite and sucrose but not from lactose. Nitrates reduced to nitrites and am- monia. Aerobic, facultative. Optimum temperature 20 to 28°C. Habitat : Sea water and marine mud. Bavendarmn (Arch. f. Mikrobiol., 3, 1932, 214) states that Pseudomonas calcis is probably synonymous with Bacterium brandti, Bacterium bauri and Bacterium feiteli described by Parlandt (Bull. Jard. imp. Bot. St. Petersburg, 11, 1911, 97-105). 56. Pseudomonas calciprecipitans Mo- lisch. (Cent. f. Bakt., II Abt., 65, 1925, 130.) From Latin, calx (calc-), lime; praecipito, to cast down headlong, to precipitate. Thin rods: 0.5 to 0.8 by 1.5 to 3.6 mi- crons, with rounded ends, often staining irregularly. Motile, with one polar flagel- lum. Gram -negative. Gelatin colonies : Circular, light brown in color (large colonies show CaCOs crystals). Gelatin stab : Surface growth with fili- form growth in depth. Liquefaction starts at bottom. Agar colonies (sea water). Grayish- white, glistening. In two to three weeks crystals of calcium carbonate form in the agar. Agar slant: Slight, whitish, surface growth, becoming thick, spreading, glis- tening, with abundant CaCOs crystals in medium. Ammonia formed. Aerobic, facultative. Optimum temperature 20°C. Habitat : Sea water. 57. Pseudomonas ichthyodermis (Wells Zobell) Zobell and Upham. (Achromo- bacter ichthyodermis Wells and Zobell, Proc. Nat. Acad. Sci., 20, 123, 1934; Zobell and Upham, Bull. Scripps Inst. Oceanography, S, 1944, 246 and 253.) From Greek, ichthys, a fish; derma, sldn. Small rods, 0.9 to 1.3 by 3 to 5 microns, occurring singly and in pairs. No spores. Encapsulated. Polar flagella. Pleomor- phic forms predominate in old cultures. Gram-negative. Requires sea water following initial isolation. The following differential me- dia are prepared with sea water. Agar colonies : Glistening, colorless, convex, circular colonies 2 to 4 mm. in diameter. Agar slants : Abundant, filiform, raised, smooth, opalescent growth. Gelatin tube : Rapid crateriform lique- faction complete in 5 days at 18°C. Sea water broth : Turbidity, with pelli- cle, little granular sediment and no odor. Milk: No growth. Casein digested when 3 per cent salt is added. Potato : No growth unless dialyzed in sea water. Then fair growth with nc pigment. FAMILY PSEUDOMONADACEAE 109 Acid from glucose, maltose, sucrose and mannitol but not from lactose or glycerol. Starch hydrolyzed. Ammonia liberated from peptone but .no hydrogen sulfide produced. Indole formed in tryptophane sea water broth. Nitrites produced from nitrates. Optimum temperature 20 to 25°C.; 30°C. incubation will kill recently isolated organisms. Aerobic, facultative. Source : Isolated from diseased kilifish (Fundulvs parvipinnis) . Habitat : Skin lesions and muscle tissue of infected marine fish. 58. Pseudomonas nigrifaciens White. (Scientific Agriculture, W, 1940, 643.) From Latin viger, black and faciens, making. Rods: 0.3 to 0.7 by 1 to 5 microns, oc- curring singly or in pairs, and having rounded ends. Actively motile, with a single polar flagellum. Gram-negative. Gelatin stab : Pigmented surface growth after 24 hours. Slight crateriform lique- faction changing to saccate. Agar colonies: Circular, convex, smooth, glistening, entire, 2 to 4 mm in diameter. Slight fluorescence in earl}- stages. The medium assumes a brownish color. Agar slant : Growth filiform, smooth, moist, glistening, with blackish pigmenta- tion at 4° and 15°C. in 48 hrs., the medium turning brownish. Slight fluorescence in early stages. Broth: Turbid after 24 hours. .A.fter 5 to 6 days a black ring and then a pellicle forms, later a black sediment. Medium turns brown. Litmus milk : A black ring appears after 3 days at 15°C. followed by a pellicle. Litmus is reduced. Alkaline reaction. No coagulation. Digested with a putrid odor. Indole not formed. Nitrites not produced from nitrates in 7 days. No gas produced. iStarch is hydroh'zed. Natural fats not hydrolyzed. Alkaline reaction produced in sucrose, maltose, lactose, glucose, mannitol and raffinose broth (pH 8.2). No gas pro- duced. Ammonia produced in peptone broth. Aerobic. Optimum pH 6.8 to 8.4. Temperature relations: Minimum 4°C. Optimum 25°C. Ma.ximum 33-35°C. Distinctive characters : No or slow growth in culture media in the absence of salt. Maximum growth and pigmenta- tion appeared with 1.5 and 2.5 per cent salt. Optimum pigmentation occurs at 4" and 15°C. Source : Several cultures isolated from samples of discolored butter. Habitat : Causes a black to reddish- brown discoloration of print butter. Evidently widely distributed in nature. 59. Pseudomonas beijerinckii Hof. (Travaux botaniques neerlandais, S2, 1935, 152.) Named for M. W. Beijerinck, Dutch bacteriologist . Small rods: Motile with polar flagella. Gelatin: No liquefaction. Indole not formed. Nitrites produced from nitrates by four out of six strains. Cellulose not decomposed. Acid from glucose. In yeast-water with 2 per cent glucose and 12 per cent NaCl no gas is produced. Pigment production : Insoluble purple pigment produced but not in all media; is localized markedly; reduced oxygen tension necessary; optimum pH 8.0; not produced in yeast-water or in peptone- water; produced only when grown in extracts of beans or some other vegetable. Aerobic. Source : Six strains isolated from beans preserved with salt. Habitat: Causes purple discoloration of salted beans. 110 MANUAL OF DETERMINATIVE BACTERIOLOGY 60. Pseudomonas salinaria Harrison and Kennedy. (Harrison and Kennedy, Trans. Royal Soc. of Canada, 16, 1922, 121; Serratia salinaria Bergey et al.. Manual, 1st ed., 1923, 93; Flavobacterium (Halobacterium) salinarium Elazari-Vol- cani. Studies on the Microflora of the Dead Sea, Thesis, Hebrew Univ., Jeru- salem, 1940, 59.) From Latin, salinae, saltworks. Probable synonym : Serratia sambhari- anus Dixit and Vachna, Current Sci., 11, 1942, 107 (see Biol. Abs., 17, 1943, 793). Halophilic growing in 30 per cent salt. From salt lake in India. Occurs as spheres and rods, 2.0 to 3.0 microns in diameter, 1.0 to 1.6 by 3.0 to 15.0 microns, occurring singly, as ovoid, amoeboid, clavate, cuneate, trun- cate, spindle, club, pear-shape, and irregular forms. Motile, frequently with a fiagellum at each pole. Gram-negative. Does not grow on ordinary culture media. Grows well on salted fish. Codfish agar (16 to 30 per cent salt) : Growth slow, smooth, raised, coarsely granular, entire, pale pink to scarlet (Ridgway chart). No acid from carbohydrate media. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 42°C. Source : Isolated from cured codfish (Harrison and Kennedy, loc. cit.). Iso- lated from salted fish by Browne (Absts. Bact., 6, 1922, 25 and Proc. Soc. Exper. Biol, and Med., 19, 1922, 321) who re- garded this pleomorphic bacterium as two organisms — Spirochaeta halophiiica and Bacterium halophiiica. Habitat: Produces reddening of dried codfish and causes rusty herring. In sea salt, and salt ponds containing not less than 16 per cent salt. 61. Pseudomonas cutirubra Lochhead. {Serratia cutirubra Lochhead, Can. Jour. of Research, 10, 1934, 275; Bacterium cutirubrum Lochhead, Jour. Bact., 27, 1934, 62; ibid., J^5, 1943, 575; Flavobac- terium {Halobacterium cutirubrum Elaz- ari-Volcani, loc. cit., 59.) From Latin, cutis, skin, hide; ruber, red. Occurs as rods and spheres. Spheres 1 to 1.5 microns in diameter. Rods 1.5 to 8.0 by 0.7-1.4 microns. Rod forms motile with single polar flagellum. Coc- coid forms motile when young. Gram- negative. No growth on ordinary media. Milk agar (20 per cent salt to satura- tion; optimum 28-32 per cent) : Colonies 3-4 mm. in diameter, round and slightly convex. Pink to rose doree (Ridgway chart). Milk agar slants: Growth filiform, slightly spreading, rather flat with smooth, glistening surface and mem- branous consistency. Proteolytic action. Liquid media: No or slight growth. Gelatin (salt ) : Pronounced liquefaction . Indole not formed, Lochhead {loc. cit.). Faint test. Gibbons (Jour. Biol. Board Canada, 3, 1936, 75). Nitrites not produced from nitrates. Diastatic action negative. No carbohydrate fermentation. Aerobic, obligate. Optimum temperature 37°C. Halophilic. Source : Isolated from salted hides. Habitat: Sea water and sea salt. 62. Pseudomonas harveyi Johnson comb. nov. {Achromobacter harveyi John- son and Shunk, Jour. Bact., 31, 1936, 587.) Named for E. N. Harvey, who studied luminous bacteria. Rods: 0.5 to 1.0 by 1.2 to 2.5 microns, occurring singly or in pairs, with rounded ends. Occasionally slightly curved; ends occasionally slightly pointed. Non-spore- forming. Capsules absent. Motile with a single polar flagellum, 2 to 3 times the length of the cell. Gram-negative. FAMILY PSEUDOMONADACEAE 111 Sea water gelatin colonies: After 24 hours at 20°C, circular, about 1.5 mm. in diameter or larger, margin slightly undu- late, sunken due to the beginning of liquefaction, interior somewhat zonate; colonies surrounded bj' a halo of numerous small secondarj' colonies, circular and finely granular. In crowded plates a large number of gas bubbles are formed. Luminescent. Sea water gelatin stab : Rapid saccate liquefaction complete in 5 days at 22°C. Abundant flocculent sediment. Sea water agar colonies : Mostly very large, 6 to 8 cm. in diameter in 24 hours, flat, highly iridescent, circular with undulate margin, or composed of narrow and close or wide filamentous growth. Occasionally small colonies appear that are circular, with entire or slightly undu- late margin, often producing irregular secondary growth, surface always smooth. Luminescent. Sea water agar slant : Growth abundant, spreading, grayishly viscous, homogene- ous, iridescent, the medium becoming rapidly alkaline when inoculated at an initial pH of 7.0. With fish decoctions added to the medium, luminescence is much brighter and growth becomes brownish after several days. Growth on autoclaved fish: Abundant, smooth, glistening, yellowish, becoming dirty brown after several days. Mild putrefactive odor. Luminescence very brilliant. Sea water containing 0.2 per cent peptone: Abundant uniform turbidity, thin pellicle, sediment accumulating over a period of several days. Luminescence at surface only unless the tube is shaken. Milk, with or without the addition of 2.8 per cent salt: Xo growth. Potato plugs resting on cotton saturated with sea water: Growth slight, somewhat spreading, slightlj^ brownish. Luminous. Indole is formed (Gore's method). Nitrites are produced from nitrates. Ammonia is produced in peptone media (Hansen method). Fixed acid from glucose, fructose, mannose, galactose, sucrose, maltose, mannitol, dextrin, glycogen, trehalose, cellobiose; slowly from salicin. Xon- fixed acid from melezitose; slight acid from sorbitol, disappearing in 24 hours. No acid from gh'cerol, xylose, arabinose, dulcitol, inositol, adonitol, erythritol, arabitol, lactose, raffinose, rhamnose, fucose or alpha methyl glucoside. Starch agar: Wide zone of hydrolysis. Hj'drogen sulfide is produced (Zobell and Fantham method). Temperature relations : Optimum 35° to 39°C. Abundant growth at 22° to 25°C. Optimum luminescence at 20° to 40°C. Not pathogenic for white rats or amphi- pods. Aerobic, facultative anaerobe. Source : Isolated from a dead amphipod (Talorchestia sp.) at Woods Hole, Massa- chusetts. Habitat : Sea water. 63. Pseudomonas phosphorescens (Fischer) Bergey et al. (Bacillus phos- phorescens Fischer, Zeitschr. f. Hyg., 2, 1887, 58; Photobacterium indicum Bei- jerinck, Arch. Neerl. d. Sci. Exactes, 23, 1889, 401 ; Bacterium phosphorescens Lehmann, Cent. f. Bakt., 5, 1889, 785; not Bacterium phosphorescens Fischer, Cent. f. Bakt., 3, 1888, 107; Bergey et al., Manual, 3rd ed., 1930, 177.) From Greek phosphoreo, to bear or bring light. See page 699 for additional synonjins. Description taken from Fischer {loc. cit.). Small, thick rods: 2 to 3 times as long as wide, with rounded ends. Motile. Stain lightly with aniline dj'es. Gelatin colonies: After 36 hours, small, circular, gray-white, punctiform. Liquefaction. Bluish to green phos- phorescence in 4 to 5 days. Blood serum: Gray-white, slimy growth. Potato: Thin white layer in 2 to 3 days. 112 MANUAL OF DETERMINATIVE BACTERIOLOGt Cooked tish: Abundant growth. En- tire surface covered with a gray -white, slimy growth. Bluish-white phos- phorescence. Alkaline broth: Slight turbidity in 24 hours. Pellicle in 3 days. Acid broth: No turbidity. No phos- phorescence. Milk: No growth. No gas formed. Not pathogenic for Iaborator.y animals. Aerobic. Optimum temperature 20° to 30°C. Source: From sea water of the West Indies. Habitat: Sea water. 64. Pseudomonas pierantonii (Zirpolo) Bergey et al. (Micrococcus pierantonii Zirpolo, Boll. del. Societa dei Natural, in Napoli, SI, 1918, 75; Cocco -bacillus pierantonii Meissner, Cent. f. Bakt., II Abt., 67, 1926, 204; Bergey et al., Manual, 3rd ed., 1930, 176.) Named for Fieran- toni, an Italian. Oval rods: 0.8 by 1.0 to 2.0 microns. Polymorphic rods, sometimes vacuolated. Motile. Gram-negative. Gelatin colonies: Circular, luminous. Gelatin stab: Not liquefied. Sepia agar colonies: Circular, white, convex, smooth, serrate edge. Intense greenish luminescence. Egg-glycerol agar slant : Yellowish- green, luminous streak. Broth: Turbid. Indole not formed. Acid from glucose and maltose, some strains also produce acid from lactose and sucrose. Best growth in alkaline media. Aerobic. Optimum temperature 33°C. Source: Isolated from the photogenic organ of the cephalopod Rondeletia minor. *65. Pseudomonas martyniae (Elliott) Stapp. (Bacterium martyniae Elliott, Jour. Agr. Res., 29, 1924, 490; Stapp, in Sorauer, Handbuch der Pflanzenkr., 2, 5 Auf., 1928, 278; Phytomonas martyniae Bergey et al.. Manual, 3rd ed., 1930, 262.) From M. L. Martynia, a generic name. Rods: 0.59 to 1.68 microns. Capsules. Chains. Motile with one to several bipolar flagella. Gram-negative. Green fluorescent pigment produced. Gelatin: Liquefied. Beef agar colonies: White, round, smooth, glistening, raised. Broth : Clouding in bands. Thin pelli- cle. Small crystals. Milk : Soft acid curd with peptoniza- tion. Nitrites produced from nitrates after 2 weeks. Indole not produced. Hydrogen sulfide production slight. Acid but not gas from glucose, galac- tose, arabinose and sucrose. No acid from rhamnose, lactose, maltose, raffinose, mannitol and glycerol. Starch hydrolysis none or feeble. Optimum temperature 26°C. Maxi- mum 37°C. Minimum 1.5°C. Optimum pH 6.0 to 6.7. pH range 5.4 to 8.9 . Aerobic. Source : Isolated from diseased leaves of the unicorn plant from Kansas. Habitat : Pathogenic on Martynia louisiana. 66. Pseudomonas striafaciens (Elli- ott) Burkholder. (Bacterium stria- faciens Elliott, Jour. Agr. Res., 35, 1927, 823; Phytomonas striafaciens Bergey et al., Manual, 3rd ed., 1930, 268; Burk- holder, Phytopath., 32, 1942, 601.) From L. stria, stripe; faciens, making. * The section covering the pseudomonads that cause plant diseases has been re- vised by Prof. Walter H. Burkholder, Cornell Univ., Ithaca, New York, April, 1943. FAMILY PSEUDOMONADACEAE 113 referring to the type of lesion caused on the blades of oats. Rods: 0.66 by 1.76 microns. Motile with one to several flagella. Capsules. Gram-negative. Green fluorescent pigment produced. Gelatin: Liquefied. Beef -peptone agar colonies : White, raised, margins entire or slightly undu- lating. Broth : Clouding in layers. Ring and slight pellicle. Milk: Alkaline, sometimes a soft curd which digests or clears. Slight production of nitrites from nitrates. Indole not produced. Acid but not gas from glucose, fructose and sucrose. No acid from lactose, mal- tose, glycerol and mannitol. Starch: Hydrolysis slight. Optimum temperature 22°C. Optimum pH 6.5 to 7.0. Aerobic. Distinctive characters: Differs from Pseudomonas coronafaciens in that the cells are somewhat smaller and the patho- gen produces a streak on oat blades instead of a halo spot. Source : Forty cultures isolated from oats gathered in various parts of America. Habitat : Pathogenic on cultivated oats, and to a slight degree, on barley. 67. Pseudomonas tomato (Okabe) comb. 710V. {Bacterium tomato Okabe, Jour. Soc. Trop. Agr. Formosa, 5, 1933, 32; Phytomonas tomato Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 422.) Named for the host plant, tomato. Probable synonym : Bacterium punc- tidans Bryan, Phytopath., 23, 1933, 897. Rods : 0.69 to 0.97 by 1.8 to 6.8 microns. Motile with 1 to 3 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Slow liquefaction. Beef -extract agar colonies: White, circular, flat and glistening. Broth: Turbid in 24 hours. Pellicle. Milk: Becomes alkaline and clears. Nitrites are usually produced from nitrates. Indole not produced. No H2S produced. Acid but not gas from glucose, sucrose and lactose. No acid from maltose and glycerol. Starch hydrolysis feeble. Slight growth in 3 per cent salt. Optimum temperature 20° to 25°C. Ma.ximum 33°C. Aerobic. Source : Isolated from diseased tomato leaves. Habitat : Pathogenic on tomato, Lyco- persicon esculentum. 68. Pseudomonas aceris (Ark) Burk- holder. {Phjitomonas aceris Ark, Phyto- path., £9, 1939, 969; Burkholder, Phyto- path., 32, 1942, 601.) From Latin acer, maple; M.L. Acer, generic name. Rods: 0.3 to 0.8 by 0.8 to 2.5 microns. Motile with 1 to 2 polar flagella. Gram- negative. Green fluorescent pigment produced. Gelatin: Liquified. Beef -extract-peptone agar : Colonies are grayish-white. Appearing in 24 hours. Broth: Turbid. Milk: Clearing with no coagulation. Nitrites not produced from nitrates. Indole not produced. Hydrogen sulfide not produced. Acid from glucose, fructose, galac- tose, arabinose, xylose, sucrose, maltose, lactose, raffinose, mannitol, glycerol and dulcitol. Slight growth in broth plus 6 per cent salt (Burkholder). Temperature : 13° to 3rC. Source : From diseased leaves of the large leaf maple, Acer macrophillum . Habitat: Causes a disease of Acer spp. 69. Pseudomonas angulata (Fromme and Murray) Holland. (Bacterium an- gulatum Fromme and Murray, Jour. Agr. Res., 16, 1919, 219; Holland, Jour. Bact., 5, 1920, 224; Phytomonas angulata Bergey 114 MANUAL OF DETERMINATIVE BACTERIOLOGY et al., Manual, 3rd ed., 1930, 267.) From L. angulatus, referring to the type of lesion produced on the tobacco leaf. Description taken from Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 24). Rods : 0.75 to 1.5 by 1.5 to 3.0 microns. Motile by 1 to 6 polar flagella. Gram- negative. Gelatin: Liquefaction. Green fluorescent pigment produced. Beef -extract agar colonies : Dull white, circular, raised, smooth and glistening. Broth: Turbid in 36 hours and greenish. Milk: Alkaline. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Lipolytic action negative (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from glucose, galac- tose, fructose, mannose, arabinose, xy- lose, sucrose and mannitol. Alkaline reac- tion from salts of citric, malic, succinic and tartaric acid. Rhamnose, maltose, lac- tose , raffinose , glycerol , sali ci n , and aceti c , lactic and formic acids are not fermented. Starch not hydrolyzed. Slight growth in broth plus 5 to 6 per cent salt (Burkholder). Facultative anaerobe. Distinctive characters: Braun (Phyto- path., 27, 1937, 283) considers this species to be identical in culture with Pseudo- monas tabaci, but they differ in the type of disease they produce. Sources: Isolated by Fromme and Murray from small angular leaf spots on tobacco. Habitat : Causes the angular leaf spot of tobacco {Nicotiana tahacum). 70. Pseudomonas aptata (Brown and Jamieson) Stevens. {Bacterium aptatum Brown and Jamieson, Jour. Agr. Res., 1, 1913, 206; Phytomonas aptata Bergey et al., Manual, 1st ed., 1923, 184; Stevens, Plant Disease Fungi, New York, 1925, 22.) From Latin aptatus adapted. Rods: 0.6 to 1.2 microns. Motile with bipolar flagella. Gram -negative. Green fluorescent pigment produced in culture. Gelatin : Liquefaction. Agar slants : Moderate growth along streak, filiform, whitish, glistening. Broth : Turbid : A pellicle formed. Milk : Becomes alkaline and clears. Nitrites not produced from nitrates. Indole not produced in 10 days. Slight amount found later. No HoS produced. Acid from glucose, galactose and sucrose. No acid from lactose, maltose and mannitol (Paine and Banfoot, Ann. Appl. Biol., 11, 1924,312). Starch not hydrolyzed. Slight growth in broth plus 7 per cent salt (Burkholder). Optimum temperature 27° to 28°C. Maximum 34° to 35°C. Minimum below 1°C. Aerobic. Source : Isolated from diseased nas- turtium leaves from Virginia and diseased beet leaves from Utah. Habitat : Pathogenic on sugar beets, nasturtiums, and lettuce. 71. Pseudomonas primulae (Ark and Gardner) Starr and Burkholder. (Phy- tomonas primulae Ark and Gardner, Phytopath., 26, 1936, 1053; Starr and Burkholder, Phytopath., 82, 1942, 601.) From L. primulus, first; M.L. Primula, a generic name. Rods: 0.51 to 0.73 by 1.0 to 3.16 microns. Motile with a polar flagellum. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar colonies : Round, convex, smooth, glistening, yellowish. Milk: Coagulated. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., S^, 1942,601). Acid but not gas from glucose, lactose, sucrose, maltose, galactose, arabinose. FAMILY PSEUDOMONADACEAE 115 glycerol, dulcitol and mannitol. Starch not hydrolyzed. Growth in broth plus 5 per cent salt. Optimum temperature 19° to 22°C. Maximum 34°C. Minimum 10°C. Optimum pH 6.8 to 7.0. Minimum 4.5 to 5.0. Facultative anaerobe. Source : Isolated from leaf-spot of Primula polyantha. Habitat : Pathogenic on Primrd'a spp. 72. Pseudomonas viridilivida (Brown) Holland. (Bacterium viridilividum Brown, Jour. Agr. Res., 4, 1915, 475; Holland, Jour. Bact., 5, 1920, 225; Phylo- monas viridilivida Bergey et al., ^Manual, 1st ed., 1923, 187.) From Latin, viridis, green; lividus, blue. Rods : 1 .0 to 1 .25 by 1 .25 to 3.0 microns. Motile with 1 to 3 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin : Slow liquefaction. Beef agar colonies: Cream white, round, smooth, translucent, edges entire. Broth: Turbid, becomes lime green. Milk: Alkaline and clears. Nitrites not produced from nitrates. Indole is produced. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid from glucose and sucrose (Burk- holder). Grows well in 4.5 per cent salt. Grows in 7 per cent salt (Burkholder). Maximum temperature 34.5°C. Mini- mum 1.5°C. Aerobic. Source : Isolated from diseased lettuce from Louisiana. Habitat: Pathogenic on lettuce, Lac- tuca saliva. 73. Pseudomonas delphinii (Smith) Stapp. (Bacillus delphini Smith, Science, 19, 1904, 417; Bacteriujii del- phinii Bryan, Jour. Agr. Res., 28, 1924, 261 ; Stapp, in Sorauer, Handbuch der Pflanzenkrankheiten, 2, 5 Auf.. 1928. 106; Phytomonas delphinii Bergey et al., Manual, 3rd ed., 1930, 261.) From Latin, delphin, a dolphin; ^LL. Del- phinium, a generic name. Rods: 0.6 to 0.8 by 1.5 to 2.0 microns. Chains present. Motile with 1 to 6 polar flagella. Capsules. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquified. Beef agar slants : Growth thin, smooth, shining, transparent, margins entire, crystals. Agar becomes dark brown. Broth : Turbid in 24 hours with delicate pellicle. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Lipolytic action negative (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid from glucose, galactose and fructose; slightly acid from sucrose. No acid from lactose, maltose, glycerol and mannitol. Starch : Hydrolysis feeble. Weak growth in broth plus 4 per cent salt. Optimum pH 6.7 to 7.1. pH range 5.6 to 8.6. Optimum temperature 25°C. Maxi- mum 30°C. Minimum 1°C. or less. Source: Isolated from black spot of delphinium. Habitat: Pathogenic on delphinium causing a black spot in the leaves. 74. Pseudomonas berberidis (Thorn- berry and Anderson) Stapp. (Phyto- monas berberidis Thornberry and Ander- son, Jour. Agr. Res., 43, 1931, 36; Stapp, Bot. Rev., 1, 1935, 407; Bacterium ber- beridis Burgwitz, Phytopathogenic Bac- teria, Leningrad, 1935, 153.) From ]\I.L. Berberis, barberry, generic name. Rods: 0.5 to 1.0 by 1.5 to 2.5 microns, occurring singly or in pairs. Motile with 2 to 4 polar flagella. Capsules present. Gram-negative (Burkholder) ; not Gram-positive as stated in original de- scription. 116 MANUAL OF DETERMINATIVE BACTERIOLOGY Green fluorescent pigment produced in culture (Burkholder). Gelatin: Not liquefied. Glucose agar slants: Growth moder- ate, filiform at first, later beaded, raised, smooth, white. Butyrous in consistency. Milk: Becomes alkaline. No other change. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., S^, 1942,601). Acid from glucose, galactose, and sucrose. Maltose and rhamnose not utilized (Burkholder). No gas from carbohydrates. Starch not hydrolyzed. Optimum temperature 18°C. Maxi- mum 30°C . Minimum 7°C . Aerobic. Sources: Repeated isolations from leaves and twigs of barberry. Habitat: Pathogenic on barberry, Ber- beris thunbergerii and B. vulgaris. 75. Pseudomonas coronafaciens (Elli- ott) Stapp. {Bacterium coronafaciens Elliott, Jour. Agr. Res., 19, 1920, 153; Phytomonas coronafaciens Bergey et al.. Manual, 1st ed., 1923, 180; Stapp, in Sorauer, Handbuch der Pflanzenkrank- heiten, 2, 5 Auf., 1928, 20.) From L. corona, halo ;/aciens, producing, referring to the lesions on oat blades. Probable synonyms: Elliott (Bact. Plant Pathogens, 1930, 122) lists as synonyms. Bacillus avenae (Russell, Johns Hopkins Univ. Thesis, 1892) and Bacillus avenae Manns and Pseudomonas avenae Manns (Ohio Agr. Exp. Sta. Bui. 210, 1909, 133; Phytomonas avenae Ber- gey et al., Manual, 3rd ed., 1930, 263). Rods: 0.65 by 2.3 microns, occurring in chains. Motile with polar flagella. Capsules. Gram-negative. Green fluorescent pigment produced in culture. Gelatin : Slow liquefaction. Nutrient agar colonies: White, becom- ing irregularly circular, flat with raised margins. Broth: Slight turbidity in 24 hours. Heavy pellicle formed. Milk: Alkaline. A soft curd formed followed by clearing. Curd sometimes absent. Nitrites not produced from nitrates. Indole not formed. No HjS formed. Not lipolytic (Starr and Burkholder, Phytopath., 3^, 1942, 601). Acid but no gas from glucose and su- crose. Starch hydrolysis slight. Slight growth in broth plus 2 per cent salt. Optimum temperature 24° to 25°C. Maximum 3rC. Minimum 1°C. Source : Numerous isolations from blighted blades of oats. Habitat : Causes a halo spot on oats (Avena sativa). Artificial inoculations show barley {Hordeum vulgar e), rye {Secale cereale) and wheat (Triticum aeslivum) to be susceptible. 75a. Pseudomonas coronafaciens var. atropurpiirea (Reddyand Godkin) Stapp. (Bacterium coronafaciens var. atropur- pureum Rcddy and Godkin, Phytopath., 13, 1923, 81 ; Stapp, in Sorauer, Handbuch der Pflanzenkrankheiten, 2, 5 Auf., 1928, 39; Phytomonas coronafaciens var. atro- purpurca Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 371.) From L. ater, black, dark; purpureus, purple, referring to the color of the lesion on brome grass. Distinctive characters : This variety differs from Pseudomonas coronafaciens in that it infects the brome-grass, Bromus inermis, where it produces a water soaked spot which is dark purple in color. Source : Numerous isolations from dis- eased brome-grass. Habitat : Pathogenic on Bromus inermis and Agropyron repens. Has been artificially inoculated on oats, Avena sativa. FAMILY PSEUDOMONADACEAE 117 76. Pseudomonas lachrymans (Smith and Bryan) Carsner. (Bacterium lachry- mans Smith and Bryan, Jour. Agr. Res., 5, 1915, 466; Carsner, Jour. Agr. Res., 15, 1918, 15; Bacillus lachrymans Holland, Jour. Bact., 5, 1920, 218; Phytomonas lachrymans Bergey et al., Manual, 1st ed., 1923, 184.) From Latin, causing tears, probably referring to the opaque drops formed on the lesion caused by this pathogen. Synonym: Elliott (Man. Bact. Plant Pathogens, 1930, 147) lists the following as a synonym: Bacillus burgeri Potebnia, Khartov Prov. Agr. Exp. Sta., 1, 1915, 37. Description from Smith and Brj^an (loc. cil.) and Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 26). Rods: 0.8 by 1 to 2 microns. Motile with 1 to 5 polar flagella. Capsules. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquefied. Beef-peptone agar colonies : Circular, smooth, glistening, transparent, whitish, entire margins. Broth: Turbid in 24 hours. White precipitate with crystals. Milk: Turns alkaline and clears. Nitrites not produced from nitrates. Indole reaction weak. No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from gUicose, fruc- tose, mannose, arabinose, .xylose, sucrose and mannitol. Alkaline reaction from salts of citric, malic and succinic acid. Maltose, rhamnose, lactose, raffinose, glycerol and salicin not fermented (Clara, loc. cit.). Starch partially digested. Not digested (Clara, loc. cit.). Growth in 3 per cent salt after 12 days. No growth in 4 per cent salt. Optimum temperature 25° to 27°C. Maximum 35°C. Minimum 1°C. Aerobic. Facultative anaerobe (Clara, loc. cit.). Source : Isolated from diseased cucum- ber leaves collected in New York, Wis- consin, Indiana and in Ontario, Canada. Habitat : Pathogenic on cucumber, Cucumis sativus, and related plants. 77. Pseudomonas maculicola (McCul- loch) Stevens. {Bacterium maculicolum McCulloch, U. S. Dept. Agr., Bur. Plant Ind. Bui., 225, 1911, 14; Stevens, The Fungi which cause Plant Diseases, 1913, 28; Phytomonas maculicola Bergey et al.. Manual, 1st ed., 1923, 189; Bacterium maccullochianum Burgwitz, Phytopatho- genic Bacteria, Leningrad, 1935, 77.) From L. maculus, spot; -cola, dweller. Rods: 0.9 by 1.5 to 3 microns. Fila- ments present. Motile with 1 to 5 polar flagella. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquefied. Beef -peptone agar colonies : Whitish, circular, shining, translucent, edges entire. Broth: Turbid. No ring or pellicle. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole production feeble. No H-S formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid from glucose, galactose, xylose, sucrose, glycerol, and mannitol. Alka- line reaction from salts of citric, malic, malonic, and succinic acid. Salicin, maltose, and salts of hippuric and tartaric not utilized (Burkholder). Slight growth in broth plus 4 per cent salt (Erw. Smith, Bact. Plant Diseases, 1920, 306). Aerobic. Optimum temperature 24° to 25°C. Maximum 29°C. Minimum 0°C. Source : Isolated from diseased cauli- flower leaves from Virginia. Habitat : Pathogenic on cauliflower and cabbage. Note: Bacterium maculicola var.japo- nicum Takimoto, Bui. Sci. Fak. Terkult 118 MANUAL or DETERMINATIVE BACTERIOLOGY Kjusu Imp. Univ., 4, 1931, 545 has not been seen. 78. Pseudomonas marginata (McCul- loch) Stapp. {Bacterium viarginatum McCulloch, Science, 5If, 1921, 115; Jour. Agr. Res., 29, 1924, 174; Phytomon'as marginata Bergey et al.. Manual, 1st ed., 1923, 188; Stapp, in Sorauer, Handbuch der Pflanzenkrankheiten, 2, 5 Auf., 1928, 56.) From L. marginatus, having a border, probably refers to the definite margin of the colony. Rods: 0.5 to 0.6 by 0.8 to 1.8 microns. Motile with 1 to 4 bipolar flagella. Cap- sules. Gram-negative. Green fluorescent pigment produced in Uschinsky's and Fermi's solutions. Gelatin: Liquefied. Agar colonies : White, circular, smooth, translucent, viscid, with definite margins at first thin but later thick and con- toured. Surface wrinkled. Milk: At first slightly acid, then alka- line. Casein digested. Nitrites not produced from nitrates. Indole production slight. Hydrogen sulfide production slight. Lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from glucose, lactose, sucrose and glycerol. Starch hydrolysis feeble. Growth in 3.5 per cent salt. No growth in 4 per cent salt. pH range, 4.6 to 9.1. Optimum temperature 30° to 32°C. Maximum 40°C. Minimum 8° to 9°C. Source : Repeatedly isolated from diseased gladiolus. Habitat: Pathogenic on Gladiolus spp. and Iris spp. 79. Pseudomonas medicaginis Sackett. (Sackett, Science, 31, 1910, 553; also Colorado Agr. Exp. Sta., Bull. 158, 1910, 11 ; Bacillus medicaginis Holland, Jour. Bact., 5, 1920, 219; Phytomonas medi- caginis Bergey et al., Manual, 1st ed., 1923, 179; Bacterium medicaginis Elliott, Bact. Plant Path., 1930, 162.) From L. medica, ancient Media; M.L. Medicago, a generic name. Rods : 0.7 by 1 .2 microns. Motile with 1 to 4 flagella. Filaments present. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Not liquefied. Nutrient agar colonies: Growth in 24 hours whitish, glistening. Broth: Turbid in 24 hours. Pellicle formed. Viscid sediment. Milk: Becomes alkaline. No change. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Starch not hydrolyzed. No gas from carbohydrates. Acid from sucrose. Slight growth in broth plus 3.75 per cent salt. Optimum temperature 28° to 30°. Maximum 37.5°C. Aerobic. Source : Isolated from brown lesions on leaves and stems of alfalfa. Habitat : Pathogenic on alfalfa, Medi- cago sp. 79a. Pseudomonas phaseolicola (Burk- holder) Dowson. {Phytomonas medi- caginis var. phaseolicola Burkholder, Phytopath., 16, 1926, 915; Bacterium medicaginis var. phaseolicola Link and Hull, Bot. Gaz., 83, 1927, 413; Pseudo- monas medicaginis var. phaseolicola Stapp and Kotte, Nachrichtenb. f. d. Deutschen Pflanzenschutzdienst, 9, 1929, 35; Dowson, Brit. Mycol. Soc. Trans., 26, 1943, 10.) From L. phaseolus, bean; M.L. Phaseolus, a generic name;-coZa, dweller. Synonym : Bacterium puerariae Hedges. (Phytopath., 17, 1927, 48 and 20, 1930, 140; Phytomonas puerariae Bergey et al.. Manual, 3rd ed., 1930, 267.) Description from Burkholder and Zaleski (Phytopath., 22, 1932, 85). FAMILY PSEUDOMONADACEAE 119 Rods : 1 by 2 microns, sometimes slightly curved, filaments present. Motile with polar flagellum. Gram- negative. Green fluorescent pigment produced in culture. Gelatin stab: Slow liquefaction. Beef extract agar: Whitish, circular colonies, 2 mm. in diameter. Edges entire. Broth: Turbid. Milk: Alkaline. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide not formed. Not lipolytic (Starr and Burkholder, Photopath., 35, 1942, 601). Acid but no gas from glucose, fructose, mannose, arabinose, xjdose, sucrose and glycerol. No acid from rhamnose, lac- tose, maltose, mannitol and salicin. Alkali from salts of citric and malic acids, but not from acetic, formic, lactic or tartaric acids. Starch and cellulose not hydrolyzed. Slight growth in broth plus 4 per cent salt. Optimum temperature 20"" to 23°C. Maximum 33°C. Minimum 2.5°C. (Hedges, loc. cit.). Optimum pH 6.7 to 7.3. Maximum S.8 to 9.2. Minimum 5.0 to 5.3. (Kotte, Phyt. Zeitsch., 2, 1930, 453.) Microaerophilic. Source : Isolated from leaves, pod and stem of beans showing halo blight. Habitat: Pathogenic on beans (Phaseo- lus vulgaris), the kudzu vine (Pueraria hirsula) and related plants. 80. Pseudomonas pisi Sackett. (Sack- ett, Colorado Agr. E.xp. Sta., Bull. 218, 1916, 19; Bacterium -pisi Erw. Smith, An Introduction to Bacterial Diseases of Plants, 1920, 474; Phytomonas pisi Ber- gey et al.. Manual, 1st ed., 1923, 181.) From Gr. pisum, the pea; M.L. Pisum, a generic name. Rods : 0.68 to 2.26 microns. Motile with a polar flagellum. Gram -negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar slants : Moderate growth in 24 hours, filiform, glistening, grayish-white. Broth: Turbid with a scum in 5 days. Milk: Alkaline, soft curd, clears. Nitrites not produced from nitrates. Indole not produced! No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but notgas from glucose, galactose and sucrose. Starch not hydrolyzed. Optimum temperature 27" to 28°C. Maximum 37.5°C. Minimum 7°C. Aerobic. Source : Ten cultures isolated from 5 collections of diseased peas showing water soaked lesions on stems and petioles. Habitat: Pathogenic on garden peas, Pisum sativum and field peas, P. sativum var. arvense. 81. Pseudomonas syringae van Hall. (Kennis der Bakter. Pfianzenziekte, Inaug. Diss., Amsterdam, 1902, 191; Bacterium syringae Erw. Smith, Bacteria in Relation to Plant Diseases, 1, 1905, 68; Phytomonas syringae Bergey et al., Manual, 3rd ed., 1930, 257.) From Latin, syringa, a nymph that was changed into a reed; M.L. Syringa, a generic name. Synonyms: Bryan (Jour. Agr. Res., 38, 1928, 225) lists Bacterium citriputeale C. O. Smith, Phytopath., 3, 1913, 69, and Bacterium citrarefaciens Lee, Jour. Agr. Res., 9, 1917, 1 {Pseudomonas citrare- faciens Stapp, in Sorauer, Handb. d. Pflanzenkrankheiten, 2, 5 Aufl., 1928, 190). Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 29) lists Bacterium vignae {Pseudomonas vignae) Gardner and Ken- drick, Science, 57, 1923, 275 {Phytomonas vignae Bergey et al., Manual, 1st ed., 1923, 188), Pseudomonas viridifaciens Tisdale and Williamson, Jour. Agr. Res., 25, 1923, 141 .{Bacterium viridifaciens Tisdale and Williams, ibid. ; Phytomonas 120 MANUAL OF DETERMINATIVE BACTERIOLOGY viridifaciens Bergey et al., Manual, 2nd ed., 1925, 208), and Phylomonas vignae var. leguminophila Burkholder, Cornell Agr. Exp. Sta. Mem. 127, 1930, 51. Wil- son (Phytopath., 30, 1940, 27) lists Phytomonas cerasi (Griffin) Bergey et al. (Pseudomonas cerasus Griffin, Science, 34, 1911, 615; Bacillus cerasus Holland, Jour. Bact., 5, 1920, 217; Bergey et al., Manual, 3rd ed., 1930, 262; Bacterium cerasi Elliott, Bact. Plant Pathogens, 1930, 109.) This would include, there- fore the following synonyms which have been listed for Phytomonas cerasi. Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 25) lists Bacterium trifoliorum Jones et al. (Jour. Agr. Res., 25, 1923, 471; Phytomonas trifoliorum Burk- holder, Phytopath., 16, 1926, 922; Pseudomonas trifoliorum Stapp, in Sor- auer, Handb. d. Pflanzenkrankheiten, 2, 5 Aufl., 1928, 177) and Bacterium hold Kendrick (Phytopath., 16, 1926, 236; Pseudomonas hold Kendrick, ibid. ; Phytomonas hold Bergey et al.. Manual, 3rd ed., 1930, 258). Wilson (Hilgardia, 10, 1936, 213) lists Pseudomonas prunicola Wormald (Ann. Appl. Biol., 17, 1930, 725), Pseudomonas cerasi var. prunicola Wilson (Hilgardia, S, 1933,83), Bacterium citripntcale C. 0. Smith (Phytopath., 4, 1913, 69; Pseudomonas citriputealis Stapp, in Sorauer, Handb. d. Pflanzen- krankheiten, 2, 5 Aufl., 1928, 190; Phyto- monas citriputealis Bergey et al., Manual, 3rd ed., 1930, 278) and Pseudomonas utiformica Clara, Science, 75, 1932, HI (Phytomonas utiformica Clara, Cornell Agr. Exp. Sta. Mem. 159, 1934, 29; Bac- terium utiformica Burgwitz, Phytopatho- genic Bacteria, Leningrad, 1935, 444). A probable synonym is Phytomonas spongiosa (Aderhold and Ruhland) Ma- grou {Bacillus spongiosus Aderhold and Ruhland, Cent. f. Bakt., II Abt., 15, 1905, 376; Pseudomonas spongiosa Braun, Die Landwirtschaft, 41, 42, 1927, 2 pp.; Bacterium spongiosum Elliott, Man. Bact. Plant Pathogens, 1930, 214; Ma- grou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 414). C. O. Smith (Phytopath., S3, 1943, 82) lists the follow- ing as a synonym : Pseudomonas hibisci (Nakata and Takimoto) Stapp (Bacteri- um hibisci Nakata and Takimoto, Ann. Phytopath. Soc. Japan, 1, 5, 1923, 18; Stapp, in Sorauer, Handbuch der Pflan- zenkrankheiten, 5 Aufl., 2, 1928, 203; Phytomonas hibisci Bergey et al., Manuals 3rd ed., 1930, 264). Description from Clara (loc. cit.). Rods: 0.75 to 1.5 by 1.5 to 3.0 microns. Motile with 1 or 2 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Beef -extract agar colonies : Circular, grayish-white with bluish tinge. Surface smooth. Edges entire or irregular. Broth: Turbid in 36 hours. No pelli- cle. Milk: Alkaline. Nitrites not produced from nitrates. Indole not produced. No HoS produced. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Slight growth in broth plus 4 per cent salt. Acid but not gas from glucose, galac- tose, mannose, arabinose, xylose, su- crose, mannitol and glycerol. Alkaline reaction from salts of citric, malic, suc- cinic and lactic acid. Rhamnose, mal- tose, lactose, raffinose, salicin, and acetic, formic and tartaric acid not fermented. Starch not hydrolyzed. Facultative anaerobe. Source : Van Hall originally isolated the pathogen from lilac. Habitat : Pathogenic on lilac, citrus, cow peas, beans, lemons, cherries and many unrelated plants. 81a. Orsini reports the following as a variety — Bacterium syringae var. capsici Orsini. (Intern. Bull. Plant Prot., 33, 1942, 33.) Pathogenic on the pepper plant (Capsicum). 82. Pseudomonas atrofaciens (McCul- loch) Stevens. (Bacterium atrofaciens FAMILY PSEUDOMONADACEAE 121 McCulloch, Jour. Agr. Res., 18, 1920, 549 ; Phytomonas atrofaciens Bergey et al., Manual, 1st ed., 1923, 185; Stevens, Plant Disease Fungi, New York, 1925, 22.) From Latin, ater, black; faciens, making, referring to the color of the lesion on wheat. Rods: 0.6 by 1 to 2.7 microns. Long chains formed in culture. Capsules present. Motile with 1 to 4 polar or bi- polar flagella. Gram -negative. Green fluorescent pigment produced in culture. Gelatin: Liquefied. Beef-peptone-agar colonies: Circular, shining, translucent, white. Broth: Growth never heavy, slight rim, and a delicate pellicle. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole: Slight production. Hj^drogen sulfide: Slight production. Acid and no gas from glucose, galac- tose and sucrose. Starch is slightly hytlrolyzed. Optimum temperature 25° to 28°C. Maximum 36° to 37°C and minimum be- low 2°C. Aerobic. Sources : Isolated from diseased wheat grains collected throughout United States and Canada. Habitat: Causes a basal glume-rot of wheat. 83. Pseudomonas cumini (Kovacev- ski) Dowson. {Phytomonas cumini Kov- acevski. Bull. Soc. Bot. Bulgarie, 7, 1936, 27; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 10.) From Greek cuminum, cumin; M.L. Cuminum, a generic name. Rods : 0.5 to 0.7 by 1 to 3 microns, oc- curring in chains and filaments. Motile with 1 to 3 polar flagella. Gram-negative. Green fluorescent pigment formed in culture. Gelatin: Rapidly liquefied. Potato agar colonies: Grayish-white, circular, glistening, smooth, butyrous. Broth: Moderate turbidity. Pseudo- zoogloea. Milk: Xot coagulated. Casein pep- tonized. Nitrites not produced from nitrates. Indole not formed. No HaS formed. Acid but not gas from glucose and sucrose. No acid from lactose or glyc- erol. Starch not hydrolyzed. ■' Temperature range 5°C to 31°C. Aerobic. Source : Isolated from blighted cumin (Cuminum). Habitat : Pathogenic on cumin and dill. 84. Pseudomonas desaiana (Burk- holder) comb. nov. (B. pyocyancus sac- charum Desai, Ind. Jour. Agr. Sci., 5, 1935, 391 ; Phytomomas desaiana Burk- holder, in Bergey et al., Manual, 5th ed., 1939, 174.) Named for Desai who first isolated the species. Rods: 0.6 to 1.2 by 1.2 to 2.2 microns. Motile with a polar flagellum. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar colonies: Grayish-blue. Raised. Broth: Light clouding. Pellicle. Milk: Peptonized without coagulation. Nitrites not produced from nitrates. Indole not formed. Glucose, sucrose, lactose and glycerol fermented without gas. Starch: Hydrolysis present. Optimum temperature 30°C. Aerobic. Source : Isolated from stinking rot of sugar cane in India and associated with a white non-pathogenic bacterium. Habitat : Pathogenic on sugar cane, Saccharu tn officinar urn . 85. Pseudomonas erodii Lewis. (Phy- topath., 4, 1914, 231; Bacterium erodii Lewis, ibid.; Phytomonas erodii Bergey et al., ^'lanual, 3rd ed., 1930, 256.) From Greek, erodius, heron; ]\I.L. Erodium, a generic name. Rods: 0.6 to 0.8 by 1.2 to 1.8 microns. 122 MANUAL OF DETERMINATIVE BACTERIOLOGY Motile with 1 to 3 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar stroke : Heavy, smooth, cream - colored growth in 24 hours. Broth: Dense clouding in 24 hours. Milk : Turns alkaline and clears, litmus reduced. Nitrites not produced from nitrates. Indole produced in 14 days. No H2S produced. Acid but not gas from glucose, sucrose, lactose and glycerol. Temperature: No growth at 35°C. Aerobic, obligate. Source: Isolations from Erodium tex- anum and 4 varieties of Pelargonium. Habitat : Causes a leaf spot of Erodium iexanum and Pelargonium spp. 86. Pseudomonas apii Jagger. (Jagger, Jour. Agr. Res., £1, 1921, 186; Phyto- monas apii Bergey et al.. Manual, 1st ed., 1923, 184; Pseudomonas jaggeri Stapp, in Sorauer, Handbuch der Pflanzenkrank- heiten, 2, 5 Aufl., 1928, 210; Bacterium jaggeri Elliott, Bacterial Plant Patho- gens, 1930, 142; Phytomonas jaggeri Magrou, in Handuroy et al., Diet. d. Bact. Path., Paris, 1937, 371.) From Latin, apium, parsley, M.L. Apium, a generic name. Description from Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 24). Rods: 0.75 to 1.5 by 1.5 to 3.0 microns. Motile with a polar flagellum. Gram- negative. Green fluorescent pigment produced in various media. Gelatin: Liquefaction. Beef -extract agar colonies: Circular, glistening, smooth, edges entire. Gray- ish-white with bluish tinge. Broth: Turbid in 36 hours. PeUicle formed. Milk: Becomes alkaline. No curd. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Acid but not gas from glucose, galac- tose, fructose, mannose, arabinose, xylose, sucrose, mannitol and glycerol. Alkaline reaction from salts of acetic, citric, malic and succinic acids. Rham- nose, maltose, lactose, raffinose, salicin, and formic, lactic and tartaric acid are not utilized. Starch not hydrolyzed. Facultative anaerobe. Distinctive characters : Pathogenicity appears limited to celery. Source : Jagger isolated this repeatedly from diseased celery leaves. Habitat: Pathogenic on celery, Apium graveolens. 87. Pseudomonas matthiolae (Briosi and Pavarino) Dowson. {Bacterium mat- thiolae Briosi and Pavarino, Atti della Reale Accad. dei Lincei Rend., 21 , 1912, 216; Phytomonas matthiolae, Bergey et al.. Manual, 3rd ed., 1930, 266; Mushin, Proc. Roy. Soc. Victoria, 53, 1941, 201; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 10.) From M.L. Matthiola, a generic name. Rods: 0.4 to 0.6 by 2 to 4 microns. Gram-positive. Gram-negative (Mush- in, loc. cit.). Green fluorescent pigment produced in culture. Gelatin : Liquefied. Beef agar colonies : White, circular colonies, slightly elevated, margins smooth. Broth: Slightly turbid. Becomes pale green. Milk: Coagulation with acid reacfon. Nitrites produced from nitrates (Mushin). Hydrogen sulfide not formed. Acid from glucose, galactose, fructose, mannose, rhamnose, glycerol, mannitol, acetic acid, citric acid, formic acid, lactic acid, malic acid, and succinic acid. Feeble acid in maltose. No acid, no gas in lactose, sucrose, raffinose, starch, salicin, and tartaric acid (Mushin). Optimimi temperature 20 to 24°C. FAMILY PSEUDOMONADACEAE 123 Maximum temperature 38.5 C. Mini- mum below 0°C. (Mushin). Limits of growth in broth are pH 4.4 to pH 9.5 (Mushin). Aerobic. Source: Isolated from vascular and parenchymatic disease of stocks, Mat- thiola incana var. annua. Habitat: Pathogenic on stocks. Note: Burkholder (Phytopath., 28, 1938, 936) and Santarelli (Rev. di Pat. Veg., 29, 1939, 364) consider this species a synonym of Pseudomonas syringae. Adam and Pugsley (Jour. Dept. Agric. Victoria, 32, 1934, 306) give a descrip- tion of a green fluorescent pathogen on stocks which is similar to Pseudomonas syringae. Mushin {loc. cit.) considers Pseudomonas matthiolae to be a distinct species. 88. Pseudomonas mors-prunorum Wormald. (Jour. Pom. and Hort. Sci., 9, 1931,251; Phytomonas mors-prunorum Wormald, Trans. Brit. Mycol. Soc, 17, 1932, 169; Bacterium mors-prunorum, ibid.) From L. mors, death; prunus, plums. Rods: Motile with a polar flagellum. Gram-positive (1931). Gram-negative (1932). Note : Possibly a green fluorescent or- ganism since it produces a faint yellow color in Uschinsky's solution. Gelatin: Liquefaction. Agar colonies : White. Broth plus 5 per cent sucrose: White and cloudy. Nitrites not produced from nitrates. Acid but not gas from glucose, lactose, sucrose and glycerol. Starch not hydrolyzed. Strict aerobe. Distinctive characters : Differs from Pseudomonas prunicola {Pseudomonas syringae) in that it produces a white cloudy growth in broth plus 5 per cent sucrose; a rapid acid production in nu- trient agar plus 5 per cent sucrose, and a faint yellow or no color in Uschinsky's solution. Source : Isolated from cankers on plum trees in England. Habitat : Pathogenic on Prunus spp. 89. Pseudomonas rimaefaciens Koning. (Chron. Bot., 4, 1938, 11 ; Meded. Phytop. Labor, Willie Comm. Scholt., 14, 1938, 24.) From L. rima, fissure; Jaciens, pro- ducing. Rods: 0.6 to 2.4 microns in length. Motile with 1 to 3 polar flagella. Gram- negative. Yellow -green fluorescent water-soluble pigment produced in culture. Gelatin : Liquified. Agar colonies: Round, convex, smooth, somewhat granular with hyaline edge. Broth : Turbid. Surface growth with a sediment in a few days. Milk: Alkaline and clears. Nitrites not produced from nitrates. Peptone, asparagin, urea, gelatin, ni- trates and ammonia salts are sources of nitrogen. Hydrogen sulfide not produced. Indole production slight. Growth with the following carbon sources plus NO3, glucose, sucrose, glycerol, succinates, malates, citrates and oxalates. Less growth with manni- tol, fructose, galactose, lactose, salicy- late. Acid is produced from the sugars. No growth with dextrin, inulin, maltose, lactose, rhamnose, salicin, tartrates, acetates, formates. Starch not hj^drolyzed. Aerobic. Optimum temperature 25°C. Maxi- mum about 37°C. Very slow growth at 14°C. Thermal death point 42° to 48°C. Source: Strains of the pathogen iso- ated from poplar cankers in France and in the Netherlands. Habitat : Pathogenic on Populus bra- bantica, P. trichocarpa and P. candicans. This may be Pseudomonas syringae since the characters are the same and both organisms can infect Impatiens sp. Pseudomonas syringae infects poplars (Elliott, Bacterial Plant Pathogens, 1930, 218). 124 MANUAL OF DETERMINATIVE BACTERIOLOGY 90. Pseudomonas papulans Rose. (Rose, Phytopath., 7, 1917, 198; Phyto- monas papulans Bergey et al.. Manual, 3rd ed., 1930, 267; Bacierivrn papulans Elliott, Bacterial Plant Pathogens, 1930, 175; Phytonionas syringae var. papulans Smith, Jour. Agr. Res., 68, 1944, 294.) From L. papulans, forming blisters. Rods : 0.6 by 0.9 to 2.3 microns. Motile with 1 to 6 polar flagella. Gram-nega- tive. Green fluorescent pigment produced in culture. Gelatin : Liquefied. Broth : Moderate turbidity in 24 hours. Milk : Alkaline and at times a soft coagulum. Nitrites not produced from nitrates. Indole: May or may not be produced. Acid but not gas formed from glucose and sucrose. Optimum temperature 25° to 28°C. Maximum 37°C. Source : Twenty-five cultures isolated from blisters on apples and from rough bark. Habitat: Pathogenic on apple trees. 91. Pseudomonas pseudozoogloeae (Honing) Stapp. (Baclcrium pseudo- zoogloeae Honing, Bull, van Het. Deli Proefstation, Medan, 1, 1914, 7; Stapp, in Sorauer, Handbuch der Pflanzen- krankheiten, 2, 5 Auf., 1928, 274; Phyto- rnonas pseudozoogloeae Bergey et al., 3rd ed., 1930, 261.) From Gr., pseudo, false; M. L. zoogloea, zooglea. Rods: 0.7 to 1.5 by 0.9 to 2.5 microns. Chains. Motile with 1 or 2 polar flagella. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar colonies: Round, flat, yellow- gray. Broth : Moderate turbidity with pseu- dozoogloea in the pellicle. Milk : Coagulation. No clearing. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide produced. Acid but not gas from glucose, lactose, maltose, sucrose and mannitol. Facultative anaerobe. Source : Isolated from the black rust of tobacco. Habitat : Pathogenic on tobacco, Nicotiana tabacum. 92. Pseudomonas tabaci (Wolf and Foster) Stevens. {Bacterium tabacum Wolf and Foster, Science, ^6, 1917, 362; also Jour. Agr. Res., 12, 1918, 449; Phyto- monas tabaci Bergey et al., Manual, 1st ed., 1923, 185; Stevens, Plant Disease Fungi, New York, 1925, 34.) From Nicotiana tabacum, tobacco. Rods : 1.2 by 3.3 microns. Motile with a polar flagellum. Gram-negative. Gelatin : Liquefaction. Potato agar colonies: Grayish-white, circular, raised, wet-shining, smooth. Milk: Alkaline; clears. Nitrites not produced from nitrates. Indole not formed. Acid from glucose, galactose, fructose, 1-arabinose, xylose, sucrose, pectin, man- nitol and glycerol (Braun, Phytopath., 27, 1937,289). Ammonium sulfate, potassium nitrate, cystine, glutamic acid, glycine, succin- imide, oxamide, acetamide, and urea can be used as nitrogen source (Braun). Starch not hydrolyzed. Aerobic. Distinctive character: Braun {loc. cit.) states that Pseudomonas tabaci and Pseu- domonas angulata are identical in culture. Source : Isolated from wildfire lesions on tobacco leaves in North Carolina. Habitat: Pathogenic on tobacco, Nico- tiana tabacum. 93. Pseudomonas lapsa (Ark) Burk- holder. (Phytomonas lapsa Ark, Phyto- path., 30, 1940, 1; Burkholder, ibid., 32, 1942, 601.) From Latin, lapsus, falling, referring to a symptom of the disease. Rods: 0.56 by 1.55 microns. Motile with 1 to 4 polar flagella. Produces fluorescence in Uschinsky's, Fermi's, and Cohn's solutions. FAMILY PSEUDOMONADACEAE 125 Gelatin: Liquefied (Burkholder). Acid but no gas is produced from glucose, sucrose, maltose, lactose, glyc- erine, arabinose, xylose, galactose, raf- finose and mannitol. Slight growth in broth plus 5 per cent salt (Burkholder). Source : Isolated from stalk rot of field corn in California; also from Diabrotica beetles. Habitat : Pathogenic on corn and sugar cane. Note : Like Pseudomonas desiana. 94. Pseudomonas bowlesiae (Lewis and \Yatson) Dowson. (Phytomonas boidesii Lewis and Watson, Phytopath.. 17, 1927, 511; Bacterimn bowlesii Elliott, Bacterial Plant Pathogens, 1930, 9G; Dowson, Trans. Brit. IMycol. Soc, 26, 1943, 9.) From 'SI. L. Boidcsia. a generic name. Rods: 0.5 to 0.7 by 1.2 to 1.6 microns, occurring singl}^ in pairs or in short chains. Motile with bipolar flagella. Gram-negative. Green fluorescent pigment produced in culture. Gelatin : Liquefied. Agar slants: Yellowish, moist, glisten- ing and viscid. Broth: Uniform turbidity- throughout. Heavy viscous sediment in old cultures. Milk: Alkaline; coagulation, with a slow peptonization. Nitrites are produced from nitrates. Indole is produced. Hydrogen sulfide is produced. Acid from glucose, maltose and xylose. No acid from sucrose. Optimum temperature 27"C. Maxi- mum 37°C. Minimum — l^C. Optimum pH 7.2. pH range 4.5 to 8.6. Aerobic. Source: Isolated from diseased, water soaked spots of bowlesia. Habitat : Pathogenic on Bowlesia sep- tentrionalis. 95. Pseudomonas intybi (Swingle) Stapp. {Phytomonas intybi Swingle, Phytopath., 15, 1925, 730; Stapp, in Sor- aurer, Handbuch der Pflanzenkrank- heiten, 2, 5 Auf., 1928, 291; Bacterium intybi Elliott, Bacterial Plant Pathogens, 1930, 142.) From Latin, intibus, endive. Description from Stapp, Cent. f. Bakt., II Abt., 9/, 1935, 232. Rods: 0.4 to 0.5 by 1.4 to 2.8 microns. Motile with one to several polar flagella. Gram-negative. Green fluorescent pigment formed in culture. Gelatin: Liquefaction. Agar colonies : White, glistening, trans- parent. Broth: Turbid with fragile pellicle, and good sediment. Milk: Coagulated. Casein not pep- tonized. Nitrites are produced from nitrates with the formation of gas. Indole not formed. Acid but not gas from arabinose, xylose and glucose. No acid from sucrose. Optimum temperature 23° to 28°C. Maximum 40° to 42°C. Minimum 0°C. Distinctive character: Differs from Psexidomonas cichorii in that it liquefies gelatin and produces nitrites from ni- trates. Source : Isolated from French endive, Cichorium inlybus by Swingle, from C. endiva and lettuce, Lactuca saliva by Stapp. Habitat : Pathogenic on endive and let- tuce, causing a rot. 96. Pseudomonas marginalis (Brown) Stevens. (Bacterium marginale Brown, Jour. Agr. Res., 13, 1918, SSQ;Phylo}nonas marginalis Bergey et al.. Manual, 1st ed., 1923, 182; Stevens, Plant Disease Fungi, New York, 1925, 30.) From Latin, margo {marginis), edge, margin; M.L. viar- ginalis, on the margin, a character of the disease. Description from Brown (loc. cit.) and Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 27). Rods : Motile with 1 to 3 polar flagella. Gram-negative. 126 MANUAL OF DETERMINATIVE BACTERIOLOGY Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar colonies: Cream-colored to yel- lowish. Brotla: Turbid, with pellicle. Milk: Alkaline. Soft curd at times. Nitrites are produced from nitrates. Not produced (Clara, loc. cit.). Indole not produced. No H2S produced. Acid but not gas from glucose, galac- tose, fructose, mannose, arabinose, xylose, rhamnose, mannitol and glycerol. Al- kali from salts of acetic, citric, malic, formic, lactic, succinic and tartaric acid. Sucrose, maltose, lactose, raffinose and salicin not fermented (Clara, loc. cit.). Starch hydrolysis feeble. None (Clara, loc. cit.). Optimum temperature 25° to 26°C. Maximum 38°C. Minimum 0°C. Aerobic. Source : Isolated from marginal lesion on lettuce from Kansas. Habitat : Pathogenic on lettuce and related plants. 97. Pseudomonas setariae (Okabe) comb. nov. (Bacterium setariae Okabe, Jour. Soc. Trop. Agr. Formosa, 6, 1934, 63; Phytomonas setariae Burkholder, in Bergey, Manual, 5th ed., 1939, -183.) From L. seia, bristle; -arius, like; M. L. Setaria, a generic name. Rods: 0.4 to 0.8 by 1.8 to 4.4 microns. Motile with a polar, seldom bipolar, flagel- lum. Gram-negative. Yellowish water-soluble pigment pro- duced in culture. Gelatin: Slow liquefaction. Beef -extract agar colonies : Circular, white, opalescent, smooth, glistening. Broth: Turbid after 18 hours. Pel- licle. Milk: Alkaline; clears. Nitrites are produced from nitrates. Indole is produced. No H2S produced. Acid but not gas from glucose, galac- tose and glycerol. No acid from lactose, maltose or sucrose. Starch: Feeble hydrolysis. Grows in 3 per cent salt. Optimum temperature 31° to 34°C. Maximum 42°C. Aerobic. Source : Isolated from brown stripe of Italian millet. Habitat: Pathogenic on Italian millet, Setaria italica. 98. Pseudomonas polycolor Clara. (Clara, Phytopath., 20, 1930, 704; Phyto- monas polycolor Clara, ibid., Bacterium polycolor Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 148.) From Gr. poly, many; L. color, color. Note: Delacroix (Comp. rend. Acad. Sci., Paris, 137, 1903, 454) describes Bacilhis aerogenosus as being a tobacco pathogen. The organism described by Delacroix might be the same as Pseudo- monas polycolor. Braun and Elrod (Jour. Bact., 4S, 1942, 40) are of the opinion that Clara's pathogen is Pseudo- monas aeruginosa. Description taken from Clara (Cornell Agr. Exp. Sta. Mem. 159, 1934, 28). Rods : 0.75 to 1 .2 by 1.05 to 3.0 microns. Motile with 1 or 2 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Beef-extract agar colonies: Gra3ash- white, circular, raised, thin transparent margins. Broth: Turbid in 36 hours with thin pellicle. Milk: Alkaline; no curd. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Lipolytic (Starr and Burkholder, Phy- topath., 5^, 1942,601). Acid but not gas from glucose, galac- tose, fructose, mannose, arabinose, xy- lose, mannitol and glycerol. Alkaline reaction from salts of acetic, citric, malic, FAMILY PSEUDOMONADACEAE 127 lactic and formic acid. Rhamnose, su- crose, maltose, lactose, raffinose and sali- cin not fermented . Starch not hydrolyzed. Facultative anaerobe. Good growth in broth plus 7 per cent salt. Optimum temperature 25" to 30°C. Maximum 37° to 39°C. Distinctive character: Differs from Pseudomonas mellea in tj'pe of lesion pro- duced, does not digest starch, nor reduce nitrates and does not form acid from lac- tose nor sucrose. Pathogenic for labora- tory animals (Elrod and Braun, Sci. 94, 1941, 520). Source : Repeatedly isolated from leaf spot of tobacco in the Philippines. Habitat: Pathogenic on tobacco. 99. Pseudomonas viridiflava (Burk- holder) Clara. {Phytomonas viridiflava Burkh., Cornell Agr. Exp. Sta. Mem. 127, 1930, 63; Clara, Science, 75, 1934, 111; Bacterium viridiflavum Burgwitz, Phyto- pathogenic Bacteria, Leningrad, 1935, 127.) From Latin viridis, green; flaviis, yellow. Description from Clara (Cornell Agr. Exp. Sta. Mem. 139, 1934, 30). Rods: 0.75 to 1.5 by 1.5 to 3.15 microns. Motile with 1 or 2 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Beef -extract agar colonies : Grayish- white, margins corrugated, edges irregu- lar. Broth: Turbid in 36 hours. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole not formed. No H2S produced. Not lipolytic (Starr and Burkholder, Phytopath., 82, 1942, 601). Acid but not gas from glucose, fruc- tose, mannose, arabinose, xylose, manni- tol and glycerol. Alkaline reaction from salts of acetic, citric, malic, lactic and succinic acids. Sucrose, lactose, mal- tose, raffinose, salicin, and salts of formic and tartaric acids not fermented. Starch: No hydrolysis. Growth in broth plus 5 per cent NaCl. Facultative anaerobe. Source: Two cultures isolated from spotted beans, one from England and one from Switzerland. Habitat : Pathogenic on bean, Phaseo- lus vidgaris. 99a. Pseudomonas viridiflava var. con- centrica (Petersen) corrib. nov. {Phyto- monas viridiflava var. concentrica Peter- sen, Tridsskr. f. Planteavl., 88, 1932, 851; Bacterium viridiflavum var. conceniricum Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 127.) From M. L. concentricus, concentric, referring to the rings on the colonies. Distinctive characters : Differs from Pseudomonas viridiflava in that it does not grow in Uschinsky's solution, and also in the shape of the colonies. Source : Isolated from the stems and leaves of blighted beans in Denmark. Habitat : Pathogenic on the bean, Phaseolus vidgaris. 100. Pseudomonas ananas Serrano. (Serrano, Philipp. Jour. Sci., 55, 1934, 355; Phytomonas ananas and Bacterium ananas Serrano, ibid, (not to be con- fused with Erwinia ananas Serrano, ibid., 86, 1928, 271); Bacterium serranoi Burgwitz, Bact. Dis. of Plants, Lenin- grad, 1936.) From Brazilian Indian, ananas, pineapple; M.L. Ananas, generic name . Rods : 0.6 by 1.8 microns. Motile with 1 to 4 polar flagella. Gram -negative. Green fluorescent pigment produced in certain media. Gelatin: Liquefied. Beef-extract glucose agar colonies : White, v^ith undulating edges, smooth to rugose, glistening to dull. Beef -extract agar: Growth scant. Broth: Feeble growth. 128 MANUAL OF DETERMINATIVE BACTERIOLOGY Milk: Becomes alkaline with curd. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Acid but not gas from glucose, xylose and mannitol. Feeble with lactose. No acid with sucrose. Starch not hydrolyzed. Optimum temperature 30° to 31°C. Maximum 45°C. Minimum 7° to 10°C. Aerobic. Habitat : Causes a rot of pineapples, Ananas comosus. 101. Pseudomonas ligustri (d'Oliveira) comb. nov. {Bacterium ligustri d'Oli- veira, Revista Agron., 24, 1936, 434.) From L. ligustrum, privet; M. L. Ligus- trum, a generic name. Rods: 0.5 to 0.7 by 1.3 to 3 microns. No chains. No capsules. Motile with 2 to 5 polar flagella. Gram-negative. Green pigment produced on Dox agar, and in broth. Gelatin: Liquefied. Beef-extract agar colonies : Growth moderate. Milky white, circular, convex. Broth: Turbid ift 24 hours. No pel- licle. Milk : Coagulated in 6 days, and later digested. Litmus slightly acid. Nitrites not produced from nitrates. Indole not produced. Ammonia not produced. No gas from carbohydrates. Acid from glucose, galactose, arabinose and man- nose. No acid from sucrose, maltose, lactose, raffinose, mannitol and salicin. Source : From diseased Japanese privet in Lisbon, Portugal. Habitat: Pathogenic on privet, Ligtis- trum japonicum. 102. Pseudomonas sesami Malkoff. (Malkoff, Cent. f. Bakt., II Abt., 16, 1906, 665; Bacterium sesami Nakata, Ann. Phyt. Soc. Japan, 2, 1930, 242; Phytomonas sesami Kovachersky, Ann. Univ. de Sofia, Fac. Agron., 8, 1930, 464.) From Gr. sesamum, sesame; M. L. Sesa- mum, a generic name. Synonym: Nakata (loc. cit.) lists Bac- ierimn sesamicola Takimoto, Jour. Plant Protect. Tokyo, 8, 1927, 433 {Phyto- monas sesamicola Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 412). Description from Nakata {loc. cit.). Rods: 0.6 to 0.8 by 1.2 to 3.8 microns. Motile with 2 to 5 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction rapid. Beef -agar colonies : Circular, flat, stri- ate, smooth, entire margins, white. Broth: Growth rapid. No pellicle. Milk: Alkaline. No coagulation. Nitrites not produced from nitrates. Indole not produced. No H2S produced. Acid but not gas from glucose. No acid from lactose, sucrose or glycerol. Starch not hydrolyzed. Optimum temperature 30°C. Maxi- mum 35°C. Minimum 0°C. Facultative anaerobe. Source : Isolated from brown spots on leaves and stems of sesame. Habitat: Pathogenic on sesame. 103. Pseudomonas tolaasii Paine. (Paine, Ann. Appl. Biol., 6, 1919, 210; Phytomonas tolaasi Bergey et al., Man ual, 3rd ed., 1930, 259; Bacterium tolaasi Elliott, Bacterial Plant Pathogens, 1930, 226.) Named for A. G. Tolaas who first reported the species. Rods: 0.4 to 0.5 by 0.9 to 1.7 microns. Motile with 1 to 5 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. , Gelatin: Liquefaction. Bouillon agar: Streak develops in 24 hours, dirty bluish-white, wet-shining and slightly raised. Broth: Turbid in 24 hours. Pellicle. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole production slight. Acid but not gas from glucose. No acid from lactose or sucrose. fa:mily pseudomoxadaceae 129 Starch hydroh-sis feeble. Optimum temperature 25°C. Source : Isolated in England from brown-spot of cultivated mushrooms. Habitat : Pathogenic on cultivated mushrooms. 104. Pseudomonas xanthochlora (Schuster) Stapp. {Bacterium xantho- chlorum Schuster, Arbeit, a. d. Kaiserl. Biolog. Anstalt. f. Land. u. Forstw., 8, 1912, 452; Phytomonas xanthochlora Ber- gey et al., Manual, 1st ed., 1923, 180; Stapp, in Sorauer, Handbuch der Pflan- zenkrankheiten, 2, 5 Auf., 1928, 213.) From Gr. xanthus, yellow; chlorus, green. Description from FJrw. Smith, Bacteria in Rel. to Plant Dis., 5, 1914, 272. Rods: 0.75 to 1.5 by 3.0 microns. Mo- tile with 1 to 3 fiagella. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Slow liquefaction. Agar colonies : Circular, slightly raised, yellow -white. Broth: Strong clouding in 24 hours. A white pellicle. Milk : Slow coagulation and clearing. Nitrites are produced from nitrates. Indole is produced after 10 days. Hydrogen sulfide produced slowlj-. Acid but not gas from glucose and galactose. Optimum temperature 27"C. Maxi- mum 44°C. Minimum 2'C. Source: Isolated from rotting potato tubers in Germany. Habitat : Pathogenic on potato tubers and a number of unrelated plants. 105. Pseudomonas rhizoctonia (Thom- as) comb. nov. {Aplanobacter rhizoctonia Thomas, Ohio Agr. Exp. Sta. Bull. 359, 1922, 211 ; Bacterium rhizoctonia Stapp, in Sorauer, Handbuch der Pflanzenkrank- heiten, 2, 5 Auf., 1928, 290; Phytomonas rhizoctonia Burkholder, Ph3'topath., 20, 1930, 7.) From Gr. rhizo, root; ctoiuis, murder. Rods: 0.5 to 0.85 by 1.4 to 1.9 microns. Non-motile. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Nutrient agar colonies: Greenish-yel- low, later olive-buff, circular, raised, slightly viscid. Broth: Turbid, pyrite yellow. Milk: Alkaline; clears. Nitrites are produced from nitrates. Indole reaction very slight. No HoS formed. Starch : Potato starch slighth- hy- drolyzed. Growth in 8 per cent salt. Optimum temperature 25° to 27''C. Maximum 38°C. Minimum 0°C. Source : Isolated from roots of lettuce showing the rosette disease. Habitat : Pathogenic on roots of let- tuce. 106. Pseudomonas barker! (Berridge) Clara. (Bacillus of pear blossom dis- ease. Barker and Grove, Ann. Appl. Biol., I, 1914, 94; Barker and Grove's organism, Doidge, Ann. Appl. Biol., 4, 1917, 50; B. barkeri Berridge, Ann. Appl. Biol., II, 1924, 73; Phytomonas barkeri Bergey et al.. Manual, 3rd ed., 1930, 265; Bac- terium barkeri Elliott, Bacterial Plant Pathogens, 1930, 95; Clara, Science, 75, 1934, 11.) Named for B. T. P. Barker who first reported the species. Description from Doidge (loc. cit.). Rods : 0.5 to 0.8 bj^ 2 to 4 microns. Motile with 1 to 4 polar flagella. Gram- negative (Burkholder), not Gram-posi- tive as stated. Green fluorescent pigment produced in culture. Gelatin: Liquefaction. Agar: Growth is white, feeble, flat, glistening, smooth edged. Broth: Slightly turbid in 24 hours. Milk: Slowly cleared. Nitrites not produced from nitrates. Indole not formed unless culture warmed. Starch slowly digested. Source : Barker made many cultures 130 MANUAL OF DETERMINATIVE BACTERIOLOGY from blighted pear blossoms. Doidge received a culture from Barker. Habitat: Causes a blossom blight of pear. 107. Pseudomonas gladioli Severini. (Severini, Annali d. Bot., Rome, 11, 1913, 420; Bacterium gladioli Elliott, Bact. Plant Pathogens, 1930, 132; Phyto- monas gladioli Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 356.) From L. gladiolus, a little sword; M. L. Gladiolus, a generic name. Rods: 0.6 by 2.3 to 2.8 microns. Mo- tile with one or more polar flagella. Gram-negative. A pale yellow water-soluble pigment found, later orange. Gelatin colonies: Cream -colored, wart- like. Rapid liquefaction. Milk: Coagulated and slowly pep- tonized. Nitrites not produced from nitrates. Indole not formed. No gas. Aerobic. Optimum temperature 28° to 30°C. Habitat : Causes a corm rot of gladiolus and other tubers. 108. Pseudomonas mellea Johnson. {Bacterium melleum Johnson, Jour. Agr. Res., ^S, 1923,489; Johnson, loc. cit., 489; Phytomonas mellea Bergey et al., Manual, 3rd ed., 1930, 254.) From L. melleus, of or belonging to honey, the color of the colonies. Rods: 0.6 by 1.8 microns. Capsules. Motile with 1 to 7 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Liquefied. Potato - glucose agar: Abundant growth, smooth, glistening, viscid, honey- colored. Broth: Turbid in 24 hours. Pellicle. Milk: Alkaline; clears. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Starch hydrolysis feeble. Growth inhibited by 4 per cent salt. Optimum temperature, 26° to 28°C. Maximum 36°C. Facultative anaerobe. Distinctive character: Differs from Pseudomonas pseudozoogloeae in that it produces on tobacco a brown instead of a black spot with a halo, is orange-yellow in culture, and turns milk alkaline. Source: Isolated from brown rusty spots on tobacco in Wisconsin. Habitat : Pathogenic on leaves of to- bacco, Nicotiana tahacum. 109. Pseudomonas betlis (Raguna- than) comb. nov. {Bacterium betle Ragunathan, Ann. Roy. Card., Pera- deniya, Ceylon, 11, 1928, 51; Aplanobac- ter betle Elliott, Bact. Plant Pathogens, 1930, 4; Phytomonas betlis Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 337.) From Malayan, belle, betel, a kind of pepper, Piper betle. Rods: 0.5 by 1.5 to 2.5 microns, occur- ring singly or in short chains. Non- motile. Gram-negative. Green pigment formed in nutrient gela- tin and in broth. Gelatin: Liquefaction. Bovril agar colonies: Honey -yellow, circular at first, later echinulate. Raised, smooth and shiny. Broth: Surface becomes cloudy in 2 days. Pellicle. No gas from lactose, maltose or sucrose. Starch is reduced. Aerobic. Source : Five cultures isolated from leaf spots on the betel vine. Habitat : Pathogenic on the betel vine, Piper betle. 110. Pseudomonas panacis (Takimoto) Dowson. {Bacterium panaxi Nakata and Takimoto, Bui. Agr. Sta. Chosen, 5, 1922, 1 ; Phytomonas panaxi Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 389; Dowson, Trans. Brit. FAMILY PSEUDOMONADACEAE 131 Mycol. Soc, 26, 1943, 10.) From Gr. panax (panicis), a plant heal-all; M. L. Panax, a generic name. Description from Elliott, Bact. Plant Pathogens, 1930, 173. Rods : 0.5 by 1.3 to 1.5 microns. Chains. Motile with 4 to 6 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: Slight liquefaction. Agar colonies : White. Milk: Coagulated. No gas from sugars. Habitat : Causes a root rot of ginseng, Panax quinquefolium. 111. Pseudomonas aleuritidis (McCul- loch and Demaree) Stapp. {Bacterium aleuritidis McCulloch and Demaree, Jour. Agr. Res., 45, 1932, 339; Stapp, Bot. Rev., 1, 1935, 408; Phytomonas aleuriti- dis Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 328.) From Gr. aleurites, of wheaten flour; M. L. Aleurites, generic name. Rods: 0.6 to 0.7 by 1.1 to 3 microns. Motile with 1 to 5 polar, rarely bipolar, flagella. Capsules present. Gram-nega- tive. Green fluorescent pigment produced in certain media. Gelatin: Not liquefied. Beef agar slants: Growth is thin, white and viscid. Broth: A heavy white surface growth in 24 hours. Sediment. Milk: Becomes alkaline, but no sep- aration. Nitrites are produced from nitrates. Indole test feebly positive. Hydrogen sulfide test feebly positive. Acid but no gas from glucose, galac- tose and glycerol. Slow acid production from sucrose, maltose and lactose. Starch hydrolysis feeble. Optimum temperature 27" to 28°C. Maximum temperature 37°C. Optimum pH 6.2 to 6.8. pH range 5.4 to 8.9. Source : Isolations from naturally in- fected tung oil trees in Georgia. Habitat: Pathogenic on the tung oil tree (Aleurites fordi), on the bean (Phaseolus vulgaris) and the castor bean (Ricinus communis). 112. Pseudomonas giycinea Coerper. (Bacterium glycineum Coerper, Jour. Agric. Research, 18, 1919, 188; Coer- per, loc. cit., 188; Phytomonas giycinea Burkholder, Phytopath., 16, 1926, 922.) From glycys, sweet; ine, like; M.L. Glycine, generic name. Synonj-m : Bacterium sojae Wolf, Phy- topath., 10, 1920, 132 (Phytomonas sojae Burkholder, Phytopath., 16, 1926, 922; Pseudomonas sojae Stapp, in Sorauer, Handb. d. Pflanzenkrankheiten, 2, 5 Aufl., 1928, 174). See Elliott, Bact. Plant Pathogens, 1930, 134; and Shunk and Wolf, Phytopath., 11, 1921, 18. Rods: 1.2 to 1.5 by 2.3 to 3 microns. Motile with polar flagella. Gram-nega- tive. Green fluorescent pigment produced in culture. Gelatin: Not liquefied. Beef-peptone agar colonies : Appear in 24 hours. Circular, creamy white, smooth, shining and convex. Margins entire. Butyrous in consistency. Milk: Litmus turns blue and later a separation of the milk occurs. Casein not digested. Nitrites not produced from nitrates. Indole test feebly positive. Not lipolytic (Starr and Burkholder, Phytopath., S2, 1942, 601). Starch not hydrolyzed. Acid from glucose and sucrose. Optimum temperature 24° to 26°C. Maximum 35°C. Minimum 2°C. Facultative anaerobe. Source : A number of cultures isolated from soy beans in Wisconsin. Habitat: Pathogenic on soybean. Gly- cine max (Soja max) . 112a. Pseudomonas giycinea var. japonica (Takimoto) comb. nov. (Bac- 132 MANUAL OF DETERMINATIVE BACTERIOLOGY terium soyae var. japonicum Takimoto, Jour. Plant Protect. Tokyo, U, 1927, 556; Bacterium glycineum var. japonicum Elliott, Bact. Plant Pathogens, 1930, 136; Phytomonas glycinea var. japonica Ma- grou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 358.) From M. L. Japonicus, Japanese. Distinctive characters : Differs slightly from Pseudomonas glycinea in size of cell, length of chains, action in milk, and color in media. Okabe (Jour. Soc. Trop. Agr., Formosa, 5, 1933, 162) gives a de- scription of the organism which leads one to believe the differences are not great enough to be varietal. Source : Isolated from leaf spots on soy bean in Formosa. Habitat : Pathogenic on soy bean, Gly- cine max. 113. Pseudomonas savastanoi (Erw Smith) Stevens. (Bacterium savastano Erw. Smith, U. S. Dept. Agr. Plant Ind Bull. 131, 1908, 31; Stevens, The Fungi which Cause Plant Diseases, 1913, 33 Phytomonas savastanoi Bergey et al.. Manual, 1st ed., 1923, 190.) Named for F. Savastano, the Italian plant path- ologist. Note: Smith (loc. cit.) lists and dis- cards the following species since they were either mixed cultures or names with no descriptions : Bacterium oleae Arcangeli , Istit. Bot. delle R. Univ. di Pisa, Ri- cerche e Lavori, fasc. 1, 1886, 109; Bacil- lus oleae tuberculosis Savastano, Atti. R. Accad. Naz. Lincei Rend. CI. Sci. Fis., Mat. e Nat., 5, 1889, 92; Bacillus pril- lieuxianus Trevisan, I generi e le specie delle Batteriacee, Milano, 1889, 19; Bacillus oleae De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 982. Description from Brown, Jour. Agr. Res., 44, 1932, 711. Rods: 0.4 to 0.8 by 1.2 to 3.3 microns. Motile with 1 to 4 polar flagella. Gram- negative. Green fluorescent pigment found in culture. Gelatin: No liquefaction. Beef agar colonies: White, smooth, flat, glistening, margins erose or entire. Broth : Turbid on the second day. No pellicle or ring. Milk: Becomes alkaline. Nitrites not produced from nitrates. No H2S produced. Acid but not gas from glucose, galac- tose and sucrose. Starch is hydrolyzed. Optimum temperature 23° to 24°C. Maximum 32°C. Minimum TC. Optimum pH 6.8 to 7.0. Maximum 8.5. Minimum 5.6. Aerobic. Source : Smith isolated his cultures from olive galls collected in California. Habitat: Pathogenic on olive. 113a. Pseudomonas savastanoi var. fraxini (Brown) Dowson. (Bacterium savastanoi var. fraxini Brown, Jour. Agr. Res., 44, 1932, 721; Phytomonas savastanoi var. fraxini Magrou, in Hau- duroy et al., Diet. d. Bact. Path., Paris, 1937, 410; Pseudomonas fraxini Skoric, Ann. Exp. For. Zagreb, 6, 1938, 66; Dow- son, Trans. Brit. Mycol. Soc, 26, 1943, 11.) From M. L. Fraxinus, a generic name. Distinctive characters: Differs but slightly from Pseudomonas savastanoi, but is pathogenic on ash and not on olive. Source : Three cultures isolated from cankers on ash. Habitat: Pathogenic on ash, Fraxinus excelsior and F. americana. 114. Pseudomonas tonelliana (Fer- raris) comb. nov. (Bacterium toncllianum Ferraris, Trattato di Patologia e Terapia Vegetale, 3rd ed., 1, 1926, 104; Phytomo- nas tonelliana Adams and Pugsley, Jour. Dept. Agr. Victoria, 32, 1934, 304.) Named for Tonelli, the Italian botanist. Synonym : Pseudomonas savastanoi var. ncrii C. O. Smith, Phytopath., 18, 1928, 503. Description from Smith (loc. cit.) un- less otherwise noted FAMILY PSEUDOMOXADACEAE 133 Rods: 0.5 to 0.6 by 1.5 to 2.5 microns. Motile with 1 to 3 polar flagella. Gram- negative (Adams and Pugsley, loc. cil.). Gelatin: No liquefaction. Potato glucose agar colonies: Flat, circular, shining, margins somewhat un- dulated. Broth: Dense clouding with partial pellicle. Milk: Alkaline. No separation. Nitrites not produced from nitrates (Adams and Pugsley). Indole produced. Not produced (Adams and Pugsley). Acid but not gas from glucose and sucrose. No acid from lactose (Adams and Pugsley). Starch not hydrolyzed (Adams and Pugsley). Distinctive character : Pseudomonas savastanoi is similar in culture but is not pathogenic on oleanders. Source: Both Ferraris and C. O. Smith isolated the pathogen from galls on olean- der. Habitat: Pathogenic on oleander, Nerium oleander. 115. Pseudomonas calendulae (Taki- moto) Dowson. {Bacleriuin calendulae Takimoto, Ann. Phytopath. Soc. Japan, 5, 1936, 341; Phytomonas calendulae Burkholder, in Manual, 5th ed., 1939, 201; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 9.) From Latin, calendae, throughout the month; M.L. Calendula, a generic name. Rods: 0.5 by 1 to 2 microns. Motile with 1 to 3 polar flagella. Gram-nega- tive. Green fluorescent pigment produced in Uschinsky's and in Cohn's solutions. Gelatin: Not liquefied. Agar colonies: Circular, smooth, fiat, dirty white. Broth: Turbid. Milk: No coagulation. Nitrites not produced from nitrates. Indole formed in small amount. No HiS produced. Acid but not gas from glucose and glycerol. No acid from lactose or sucrose. Starch not hydrolyzed. Optimum temperature 27° to 30°C. Maximum 37°C. Minimum 0° to 7°C. Habitat : Pathogenic on marigolds. Calendula officinalis. 116. Pseudomonas cichorii (Swingle) Stapp. (Phytomonas cichorii Swingle, Phytopath., 15, 1925, 730; Stapp, in Sor- auer, Handbuchder Pflanzenkrankheiten, 2, 5 Auf., 1928, 291; Bacterium cichorii Elliott, Bact. Plant Pathogens, 1930, 112.) From Gr. cichoria, chicory; M. L. Cichoriiim, generic name. Probable synonyms : Pseudomonas en- diviae Kotte, Phyt. Ztschr., 1, 1930, 609; Phytomonas endiviae (Kotte) Clara, Cornell Agr. Exp. Sta. Mem. 159, 1934, 26; and Bacterium formosaniim Okabe, Jour. Soc. Trop. Agr., Formosa, 7, 1935, 65. Description from Clara (loc. cit.) which is a description of a culture of Pseudo- monas endiviae from Kotte. Swingle's description is verj^ meager. Rods : 0.75 to 1.5 by 1.5 to 3.75 microns. Motile with 1 or 2 polar flagella. Gram- negative. Green fluorescent pigment produced in culture. Gelatin: No liquefaction. Beef -extract agar colonies : Circular, grayish-white with bluish tinge, raised with slightly irregular edges. Broth : Turbid in 36 hours with a smooth viscous pellicle. Milk: Alkaline. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from glucose, galac- tose, fructose, mannose, arabinose, xy- lose, mannitol and glycerol. Alkaline production from salts of acetic, citric, lactic, malic, succinic and tartaric acids. 134 MANUAL OF DETERMINATIVE BACTERIOLOGY Rhamnose, maltose, sucrose, lactose, raf- finose and salicin not utilized. Starch not hydrolyzed. Slight growth in broth plus 6 per cent NaCl. Optimum pH 6.8 to 7.1. Maximum 9.2 to 9.4. Minimum 5.0 to 5.3 (Kotte, Phyt. Ztsch., 2, 1930, 453). Facultative anaerobe. Distinctive characters: Differs from Pseudomonas inlyhi in that it does not liquefy gelatin or reduce nitrates to ni- trites. Source : Isolated from rot of French endive, Cichorium intybus by Swingle and by Okabe, and from C. endivia by Kotte. Habitat : Pathogenic on endive, lettuce and larkspur. 117. Pseudomonas cissicola (Taki- nioto) comb. nov. {Aplanobacter cissicola Takimoto, Ann. Phytopath. Soc. Japan., 9, 1939, 43.) From Greek, cissus, ivy; M. L. Cissus a generic name. Rods: 0.5 to 0.9 by 1.0 to 2.0 microns. Non-motile. Capsules. Gram-negative. Green fluorescent pigment formed in Uschinsky's solution. Gelatin: No liquefaction. Potato-extract agar colonies: Circular, convex, smooth, and dirty white. Broth: Feeble clouding followed by precipitation of pellicle and rim. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide not produced. No acid nor gas from sucrose, glucose, lactose and glycerol. Starch is not digested. Salt toleration is 3 per cent. Optimum temperature 30°C. Maxi- mum 35°C. Minimum lO'C. Thermal death point 49° to 50°C. Source : Isolated from black spots on leaves of Japanese ivy, Cissus japonica in Japan. Habitat : Pathogenic only on Cisstis japonica. 118. Pseudomonas nectarophila (Doidge) Rosen and Bleeker. {Bac- terium nectarophila Doidge, Ann. Appl. Biol., 4, 1917, 73; Phytomonas nectaro- phila Bergey et al., Manual, 3rd ed., 1930, 262 ; Rosen and Bleeker, Jour. Agr. Res., It6, 1933, 98.) From Gr. nectar, nectar; philus, loving. Rods: 0.5 to 0.7 by 0.6 to 1.5 microns. Motile with 1 to 5 polar flagella. Cap- sules. Gram-negative. Green fluorescent pigment produced in culture. Gelatin: No liquefaction. Nutrient agar colonies : Yellowish- white, wet-shining, smooth, margins ir- regular. Broth: Heavy turbidity in 24 hours. Sediment. Milk: Cleared. Nitrites not produced from nitrates. Indole not formed. Acid from glucose and galactose. No acid from sucrose. Starch hydrolysis feeble. Optimum temperature 25 to 30°C. Facultative anaerobe. Distinctive character: Differs from Pseudomonas barkeri in that it does not liquefy gelatin, nor produce indole. Pro- duces capsules. Source : Isolated from blighted pear blossoms in South Africa. Habitat: Pathogenic on pear blossoms. 119. Pseudomonas viburni (Thorn- berry and Anderson) Stapp. {Phytomonas viburni Thornberry and Anderson, Phy- topath., 21, 1931, 912; Stapp, Bot. Rev., 1, 1935, 407; Bacterium viburni Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 160.) From L. viburnum, the way- faring tree; M. L. Viburnum, a generic name. Rods: 0.5 to 1.0 by 1 to 2.0 microns. Capsules present. Motile with 2 to 4 polar flagella. Gram-negative (Burk- holder) ; not Gram-positive as stated. Green fluorescent pigment produced in culture (Burkholder). Gelatin: No liquefaction. Glucose beef -extract colonies: Dull gray, circular, edges entire. FAMILY PSEUDOMONADACEAE 135 Broth: Turbid with pellicle. Milk: Alkaline. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid from glucose and galactose, but not sucrose (Burkholder). Starch : No hydrolysis. Slight growth in 3.5 per cent salt (Burkholder). Optimum temperature 25°C. Mini- mum 12°C. Maximum 35°C. Aerobic. Source : Isolated from angular leaf spots and stem lesions on arrow-wood, Viburnujn opulus, etc. Habitat : Pathogenic on Viburnum spp. 120. Pseudomonas mori (Bo3^er and Lambert) Stevens. {Bacterium vinri Boyer and Lambert, Compt. rend. Acad. Sci. Paris, 117, 1893, 342; Bacterium mori Boyer and Lambert emend. Erw. Smith, Science, 31, 1910, 792; Stevens, The Fungi which Cause Plant Diseases, 1913, 30; Bacillus mori Holland, Jour. Bact., 5, 1920, 222; Phytomonas mori Bergey et al., Manual, 1st ed., 1923, 191.) From Gr. moriun, mulberry; ]\L L. Moms, a generic name. Synonyms: Elliott (Bact. Plant Path- ogens, 1930, 166) lists Bacillus cuboni- anus Macchiati, Staz. Sperim. Agr. Ital., S3, 1892, 228 (Macchiati described the disease due to Pseudomonas mori, but gave an incorrect description of the patho- gen) ; also Bacterium cubonianum Fer- raris, Curiano le Plante, 6, 1928, 180 (Ferraris uses Macchiati 's name but the description of Pseudomonas ynori). Description from Smith {loc. cit.). Rods: 0.9 to 1.3 by 1.8 to 4.5 microns. Motile with a polar flagellum. Gram- negative. Green fluorescent pigment produced in culture. Gelatin : Not liquefied. Agar colonies : White, slow -growing, smooth, flat, edges entire becoming un- dulate. Milk: Becomes alkaline and clears. Nitrites not produced from nitrates. Indole none or feeble production. Hydrogen sulfide not produced (Okabe, Jour. Soc. Trop. Agr., 5, 1933, 166). No growth in broth plus 4 per cent salt (Okabe, loc. cit.). No gas from carbohydrates. Temperature range 1°C to 35°C. Source: Smith isolated the pathogen from blighted shoots of mulberry from Georgia. Also received cultures from Arkansas and the Pacific Coast. Habitat : Pathogenic on mulberry, Morus. 121. Pseudomonas stizolobii (Wolf) comb. nov. {Aplanobacter stizolobii Wolf, Phytopath., 10, 1920, 79; Bacterium stiz- olobii McCulloch, Phytopath., 18, 1928, 460; Phytomonas stizolobii Bergey et al.. Manual, 3rd ed., 1930, 280.) From Gr. siizo, to prick; lobium, a little lobe; Stizolobium, a generic name. Rods: 0.6 to 0.7 by 1.0 to 1.6 microns. Non-motile (Wolf). Motile with a short polar flagellum (McCulloch). Capsules. Gram-negative. Gelatin : No liquefaction. Agar colonies : Circular, smooth, white, raised and opaque. Margins entire to slightly undulate. Broth: Slightly turbid throughout. No pellicle or ring. Milk: Alkaline. Nitrites not produced from nitrates. Indole not formed. No acid or gas in peptone broth plus sugars. Starch not hydrolyzed. Optimum temperature 25° to 28°C. Distinctive characters : Differs from Pseudomonas sojae (Pseudomonas gly- cinea) in the smaller size of cell, and absence of pellicle and dense clouding of broth. The pathogen does not infect soy bean. Source : Isolated from the leaf spot of velvet bean. Habitat : Pathogenic on velvet bean, Stizolobium deeringianium. 136 MANUAL OF DETERMINATIVE BACTERIOLOGY 122. Pseudomonas viciae Uyeda. (Uyeda in Takimoto, Jour. Plant Pro- tect., Japan, 2, 1915, 845; Bacterium vi- ciae Nakata, see Elliott, Bact. Plant Pathogens, 1930, 259; Phytomonas viciae Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 430.) From L. vicia, vetch; M. L. Vicia, a generic name. Rods: 0.5 to 0.8 by 1.2 to 2.0 microns. Motile with 2 to 4 polar flagella. Gram- positive. Green fluorescent pigment produced in culture. Gelatin colonies : Pale white, glisten- ing, finally turning brown. No liquefac- tion. Milk : Coagulates and clears. Nitrites not produced from nitrates. No H2S produced. Facultative anaerobe. Habitat : Pathogenic on the broad bean {Vicia faba), the turnip (Brassica rapa), the carrot (Daucus carota) and the sweet potato {Ipomoea batatas). 123. Pseudomonas alliicola Burk- holder. (Burkholder, Phytopath., 32, 1942, 146 ; Phytomonas alliicola Burk- holder, ibid.) From L. allium, onion; -cola, dweller. Rods: 0.7 to 1.4 by 1.05 to 2.8 microns. Motile with 1 to several polar flagella, at times bi -polar. Gram-negative. Gelatin : Liquefaction. Beef -extract peptone agar streaks : Moderate in growth, white at first, later dirty in appearance, edges wavy, consist- ency viscid. Medium deep brown. Potato-glucose agar frequently be- comes greenish. Broth: Turbid with light pellicle. Brown. Milk : Cleared and litmus reduced . Neutral. Nitrites produced from nitrates. Indole not produced. Hydrogen sulfide not produced. Lipolytic action very strong. Acid but no gas from 1-arabinose, d- xjdose, rhamnose, glucose, d-galactose, fructose, d-lactose, maltose, sucrose, glycerol, mannitol and salicin. Alkali from salts of acetic, citric, formic, hip- puric, lactic, malic, succinic, tartaric acids. Starch not hydrolysed. Slight growth in broth plus 4 per cent salt. Aerobic. Optimum temperature 30°C. Maxi- mum 41°C. Minimum 5°C. Source: Seven isolates from storage rot of onion bulbs. Habitat: Pathogenic on onion bulbs, Allium cepa. 124. Pseudomonas gardeniae Burk- holder and Pirone. (Burkholder and Pirone, Phytopath., 31, 1941, 194; Phyto- monas gardeniae Burkholder and Pirone, ibid.) From M. L. Gardenia, a generic name. Rods: 0.75 by 2.4 microns. Motile with 1 to 2 polar flagella. Gram-nega- tive. Gelatin: Liquefaction. Beef -extract peptone agar colonies: Growth fair, white to dirty gray and vis- cid. Medium becoming dark brown. Potato-glucose agar : No brown color. Broth: Turbid with pellicle. Dark brown. Milk: Soft curd with pellicle. Clears in zones. Litmus reduced. Nitrites produced from nitrates. Hydrogen sulfide not produced. Indole not formed. Acid from glucose, galactose, xylose, rhamnose, sucrose, maltose, mannitol, glycerol, and salicin. Alkali produced from the salts of citric, malic, malonic, succinic, tartaric and hyppuric acids. Good growth in tyrosine and in aspara- gine broth. Starch is not hydrolyzed. Aerobic. Source: Eight isolates from leaf spots of gardenias in New Jersey. Habitat: Pathogenic on leaves of Gar- denia jasminoides. 125 Pseudomonas caryophylli Burk- holder. (Burkholder, Phytopath., 31, FAMILY PSEUDOMOXADACEAE 137 1941, 143; Phytomonas caryophylli, Burkholder, ibid.) From M. L. Caryo- phyllus, an old generic name. Rods: 0.35 to 0.95 by 1.05 to 3.18 mi- crons. At times slightly curved. Motile with 1 to several polar flagella. Fre- quently bipolar. Gram-negative. Gelatin: Liquefaction after 3 to 4 weeks. Potato glucose agar colonies: 3 to 4 mm in diameter, circular, smooth, glis- tening, edges entire. Color is tan to gray mauve. Old culture dark brown. Con- sistency butj^rous. Broth: Turbid with a white sediment. Milk: Litmus slowly becomes blue. Slight reduction at bottom of tube. No clearing. Nitrites produced from nitrates. Also ammonia and gas are produced in a synthetic nitrate medium. Asparagine, KXO3 and XH4H2PO4 can be utilized. Indole not formed. Hydrogen sulfide not formed. Lipolytic action slight to moderate. Acid from 1-arabino.se, d-xylose, rham- nose, glucose, d-galactose, fructose, d- lactose, maltose, and sucrose, glj'cerol, mannitol, and salicin. Alkali with so- dium salts of acetic, citric, formic, hip- puric, lactic, malic, maleic, succinic and tartaric acid. Starch not hydrolyzed. Aerobic. Optimum temperature 30° to 33°C. Maximum 46°C. Minimum 5^C. or less. Slight growth in broth plus 3.5 per cent salt. Source: Isolated first by L. K. Jones and later by W. H. Burkholder from dying carnation plants from Spokane, Washington. Twelve isolates used in description. Habitat : Pathogenic on roots and stalks of the carnation, Dianthus caryo- phylliis. 126. Pseudomonas solanacearum Erw. Smith. (Bacillus solanacearum Erw. Smith, U. S. Dept. Agr., Div. Veg. Phys. and Path., Bui. 12, 1896, 10; Bacterium solanacearum Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 73; Erw. Smith, Bacteria in Relation to Plant Diseases, S, 1914, 178; Phytomonas so- lanacearum Bergej' et al., ^Manual, 1st ed., 1923, 186.) From L. solanum night- shade; AI. L. Solanaceae, a plant famih'. Probable synonyms: Elliott (Bact. Plant Pathogens, 1930, 203) lists the following: Bacillus nicotianae Uyeda, Cent. f. Bakt., II Abt., 13, 1904, 327; Bacillus sesami Malkoff and Pseudo- monas sesami Malkoff, Cent. f. Bakt., II Abt., 16, 1906, 664; Bacillus rnusae Rorer, Phytopath., 1, 1911, 45; Bacillus tnusaru?n Zeman, Rev. Facul. Agr. L'niv., La Plat, 14. 1921, 17; Erwinia nicotianae Bergey et al.. Manual, 1st ed., 1923, 172; Phytomonas ricini Archibald, Trop. Agr., Trinidad, 4, 1927, 124. Description taken from Elliott {loc. cit.). Rods: 0.5 to 1.5 microns. Motile with a polar flagellum. Gram-negative. Gelatin: Xakata (Jour. Sci. Agr. Soc. Tokyo, 294, 1927, 216) states there are two forms, one of which shows slight liquefaction. The other shows no lique- faction. Agar colonies: Small, irregular, round- ish, smooth, wet-shining, opalescent, becoming brown. Broth: Slight pellicle. Broth turns brown . ]Milk : Cleared without precipitation of casein. Nitrites produced from nitrates. Indole not formed. Hydrogen sulfide not produced (Burk- holder). Glucose, sucrose, glycerol, sodium citrate, peptone, tyrosine, asparagine and glutanic acid are utilized (Mushin, Aus- tral. Jour. Expt. Biol, and Med., 16, 1938, 325). Nitrogen sources utilized are ammonia, nitrates (KNO3) asparagine, tyrosine, peptone and glutamic acid, but not potas- sium nitrite (Mushin, loc. cit.). Starch not hydrolyzed. 138 MANUAL OF DETERMINATIVE BACTERIOLOGY Optimum temperature 35° to 37°C. Maximum 41 °C. Minimum 10°C. Pathogenicity readily lost in culture. Source : Isolated from brown-rot of solanaceous plants. Habitat : Soil pathogen in warm moist climates attacking numerous species of plants, especially potato, tobacco, and tomato. . 126a. Pseudomonas solanacearum var. asiatica (Erw. Smith) Stapp. {Bac- terium solanacearum var. asiaticum Erw. Smith, Bact. in Relation to Plant Dis- eases, 3, 1914, 282; Stapp, in Sorauer, Handbuch der Pflanzenkrankheiten, 2, 5 Auf., 1928, 253; Phytomonas solana- cearum var. asiatica Magrou, in Hau- duroy et al., Diet. d. Bact. Path., Paris, 1937, 414.) From Gr. asiaticus, asiatic. Distinctive characters: Differs from Pseudomonas solanacearum in that it turns litmus milk and cream red. Source : Isolated by J. A. Honing from diseased tobacco plants in Medan, Suma- tra. 127. Pseudomonas castaneae (Kawa- mura) comb. nov. {Bacterium castaneae Kawamura, Ann. Phytopath. Soc. Japan, S, 1934, 15; Phytomonas castaneae Ma- grou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 343.) From M. L. Castanea, a generic name. Rods: 0.8 to 1.2 by 1.0 to 1.8 microns. Motile with 1 to 5 polar flagel la. Gram- negative . Gelatin: Liquefied. Beef agar colonies : White, circular, edges slightly undulate, viscid. Milk: No coagulation. Peptonized. Acid but not gas from glucose, sucrose and glycerol. No acid from lactose. Optimum temperature 25° to 27°C. Maximum 35°C. Minimum 3°C. Facultative anaerobe. Habitat : Causes water-soaked spotting on leaves and shoots of chestnut, Cas- tanea. 128. Pseudomonas seminum Cayley. (Cayley, Jour. Agr. Sci., 8, 1917, 461; Bacterium seminum Stevenson, Foreign Plant Dis., U.S.D.A. Office of Sec'y-, 1926, 141 ; Phytomonas seminum Bergey et al., Manual, 3rd ed., 1930, 272.) From L. semen {seminus) seed. Rods : 1 .0 by 4.0 to 5.0 microns. Spore- like bodies present. Capsules. Motile with a single flagellum. Gram-positive. Gelatin: Rapid liquefaction. Agar colonies: White, more or less circular, transparent, spreading. Broth: Turbid. Pellicle. Litmus milk: Milk becomes clear and apricot color. Nitrites produced from nitrates. Acid but not gas from glucose and sucrose. No acid from lactose. Starch : No hydrolysis. Optimum temperature 25°C. Facultative anaerobe. Source : Isolated from seeds, stems and pods of diseased peas in England. Habitat : Pathogenic on peas. 129. Pseudomonas passiflorae (Reid) comb. nov. {Phytomonas passiflorae Reid, New Zealand Jour. Sci. and Tech., 22, 1939, 264a.) From L. passio, passion; flora, flower; M. L. Passiflora, a generic name. Rods: 0.2 to 0.5 by 1.2 to 3.2 microns. Motile with 1 to 5 polar flagella. Cap- sules present. Gram-negative. Gelatin : Liquefied. Beef-peptone agar colonies : Small, flat, smooth, dry, shining, translucent, gray- ish and butyrous. Broth : Turbid in 4 days. Transient pellicle. Milk: Slightly alkaline. No coagula- tion nor clearing. Nitrites not produced from nitrates. No growth on synthetic nitrate agar. Indole not formed. Hydrogen sulfide not formed. Acid reaction occurs in galactose, starch and sucrose. No gas. Starch is not hydrolysed. Source : From diseased leaves and fruit of the passion-fruit in New Zealand. Habitat : Pathogenic on Passiflora edulis. FAMILY PSEUDOMONADACEAE 139 130. Pseudomonas fabae (Yu) comb, nov. {Phytomonas fabac Yu, Bull, of the Chinese Bot. Soc, 2, 1936, 34.) From Latin, faba bean. Rods: 0.8 to 1.1 by 1.1 to 2.8 microns. Motile with 1 to 4 polar flagella. Gram- negative. Gelatin: Liquefied. Nutrient agar colonies : Circular, en- tire, viscid, glistening, raised, smooth to wrinkled, white to salmon. Medium amber. Broth: Turbid after 12 hours. Pellicle. Milk : Growth slow. Clears. Nitrites produced from nitrates. Indole production slight. Hydrogen sulfide not produced. Acid but not gas from glucose. No acid nor gas developed from arabinose, xylose, fructose, galactose, sucrose, lactose, maltose, raffinose, dextrin, in- ulin, mannitol or adonitol in a 1 per cent Bacto-peptone broth. Starch: Verj^ weak diastatic action. Optimum temperature 35°C. ^laxi- mum 37 to 38"C. Minimum 4°C. Ther- mal death point 52 to 53°C. Aerobic. Growth retarded in 2 per cent salt. Very slight growth in 3 per cent salt. Source: From diseased broad beans at Nanking, China- Habitat : Pathogenic on broad or Wind- sor bean, Vicia faba. 131. Pseudomonas astragali (Taki- moto) comb. nov. {Bacterium astragali Takimoto, Jour. Plant Protect., 17, 1930, 732; Phytomonas astragali Burkholder, in Manual, 5th ed., 1939, 197.) From M. L. Astragalus, a generic name. Description translated by Dr. K. Togashi. Rods: 0.7 to 0.8 by 1.2 to 2.2 microns. Motile, with 1 or 2 flagella. Gram- negative. Gelatin: Liquefied. Agar plates: Growth somewhat slow, colorless or grayish-white, entire margins, more or less aqueous, butyrous. Uschinsky's medium : Growth vigorous, turbid, not viscid, ring, and sediment. Milk: No coagulation of casein, slow digestion. Alkaline. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide produced in small amount. No acid or gas from glucose, sucrose, lactose and glycerol in broth. Starch not hydrolyzed. Temperature relations : Minimum be- low 5° and maximum 33'^C. Thermal death point 50° to 51 °C. Aerobic. Source : Species isolated from Astra- galus sp. Habitat : Causes a black leaf -spot of Astragahis sp. 132. Pseudomonas colurnae (Thorn- berry and Anderson) comb. nov. {Phyto- monas colurnae Thornberry and Ander- son, Phytopath., 27, 1937, 948.) From the species, Corylus colurna. Rods: 0.8 to 1.0 by 1.0 to 1.8 microns. Single, in pairs or chains. Capsules. Motile with 1 to 2 polar flagella. Gram- negative. Gelatin: Liquified. Glucose agar slants: Growth filiform, raised, dull, smooth, opaque and viscid. Broth : Moderate turbidity. Ring. Milk : Peptonization complete with acid production. No reduction of litmus nor coagulation. Nitrites not produced from nitrates. Indole not produced. Hydrogen sulfide not produced. No appreciable amount of gas from xylose, glucose, sucrose or glycerol. Starch hydrolyzed. Optimum temperature 21°C. Mini- mum 5°C. Maximum 35°C. Thermal death point oO^C. Aerobic. Source : From leaves and young stems of the Turkish hazelnut in Illinois. Habitat : Pathogenic on the Turkish hazelnut, Coryltis colurna. 140 MANUAL OF DETERMINATIVE BACTERIOLOGY 133. Pseudomonas maublancii (Foex and Lansade) comb. nov. (Bacterium maublancii Foex and Lansade, Comp. rend. Acad. Sci. Paris, 202, 1936, 2174; Phytomonas maublancii Burkholder, in Manual, 5th ed., 1939, 198.) Named for M. Maublanc, French colonial plant pathologist. Rods : 0.4 by 1.3 microns. Motile with 1 to 3 polar flagella. Gram-negative. Gelatin: Liquefied. Gelatin colonies: Round, translucent, margins entire. Broth: Thin pellicle. Milk: Not coagulated; clears. Nitrites not produced from nitrates. Indole not formed. No HoS formed. Carbohydrates not fermented. Ammonia produced. Growth in Fermi's solution, not in Us- chinsky's solution. Source : Isolated from rotting vascular and parenchymatic tissue of banana stalks. Habitat: Causes a disease of the banana plant. 134. Pseudomonas polygon! (Thorn- berry and Anderson) coynb. nov. (Phyto- monas polygoni Thornberry and Ander- son, Phytopath., 27, 1937, 947.) From Gr. polygonum, knot-weed; M. L. Poly- gonum, a generic name. Rods: 0.5 to 1.5 by 1.5 to 2.5 microns. Motile with 2 to 8 bi -polar flagella. Cap- sules. Gram-positive (?). Other spe- cies reported by these investigators as Gram-positive have proved to be Gram- negative on a retest (Burkholder). Gelatin: Liquified. Brown. Glucose agar slant: Abundant, fili- form, flat, dull, smooth, pale olive-gray, butyrous. Medium turns brown. Broth: Turbid. Pellicle. Milk: Alkaline and clears. Litmus not reduced. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide not produced. No appreciable amount of gas from car- bohydrates. Starch : No hydrolysis. Optimum temperatures 18°C. Mini- mum 7°C. Maximum 35°C. Aerobic. Source : From diseased leaves of Poly- gonum convolvulus in Illinois. Habitat : Pathogenic on black bind- weed, Polygonum convolvulus. 135. Pseudomonas iridicola (Taki- moto) Stapp. (Bacterium iridicola Taki- moto. Fungi, Nippon Fungological Soc, 1, 1931, 24; Stapp, Bot. Rev., 1, 1935, 408; Phytomonas iridicola Burkholder, in Manual, 5th ed., 1939, 198.) From Gr. iris (iridis) , iris, a rainbow; -cola, dweller. Rods: 0.7 to 0.8 by 1.2 to 2 microns. Motile with 1 to 3 polar flagella. Gram- negative. Gelatin: Liquefied. Beef agar colonies: White, circular, raised or convex. Milk: Clears without coagulation. No acid or gas from carbohydrates. Starch digested. Optimum temperature 38°C. Mini- mum 4°C. Source : Isolated from a brown leaf spot of iris. Habitat : Pathogenic on Iris tectorum and Iris japonica. 136. Pseudomonas levistici Oster- walder. (Ostcrwalder, Cent. f. Bakt., II Abt., 25, 1909, 260; Bacterium levistici Stevenson, Foreign Plant Dis., U. S. Dept. Agr., Oflfice of Sec'y-, 1926, 101; Phytomonas levistici Magrou, in Hau- duroy et al.. Diet. d. Bact. Path., Paris, 1937, 373.) From M. L. Levisticum, a generic name. Rods: 0.5 to 0.7 by 1.1 to 1.5 microns. Motile with a polar flagellum. Gram- negative. Gelatin: Colonies greenish-white. Liquefaction. Nutrient agar: Good growth at room temperature. Yellowish-white. FAMILY PSEUDOMONADACEAE 141 Broth: Pellicle. Indole formed. No H-iS produced. Source: Isolated from spots on the leaves of lovage. Habitat: Pathogenic on lovage, Levis- ticum officinale. 137. Pseudomonas radiciperda (Jav- oronkova) Stapp. {Bacterium radici- perda Javoronkova, Bull. Plant Protect., Leningrad, Ser. II, .5, no. 1, 1932, 161; Stapp, Bot. Rev., 1, 1935, 408; Phyto- monas radiciperda Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 401.) From L. radix (radicis), root; pcrdo, to destroy. Description from Javoi'onkova, Rev. App. Myc, 11, 1932, 652. Rods : O.S by 1 to 2 microns. Capsules. Motile with 1 or 2 polar flagella. Gram- negative. Gelatin: Liquefaction. Beef-peptone agar colonies: Round, smooth, shining, white to pale yellow. Milk: Peptonized. Indole not formed. No H-jS formed. Acid but not gas from carbohydrates. Optimum temperature 23° to 25°C. Aerobic. Habitat : Causes a root rot of red clover (Trifolium pratense), lentils (Lens escu- lenta) and lucerne. 138. Pseudomonas melophthora Allen and Riker. (Allen and Riker, Phyto- path., 22, 1932, 557; Bacterium meloph- thorum Allen and Riker, ibid.; Phyfo- monas melophthora Allen and Riker, ibid.) From Gr. melum, apple; phthora, destroyer. Rods: 0.68 by 1.32 microns. Motile with 2 polar flagella. Gram-negative; Gram-positive cells appear in old cultures. Gelatin : No liquefaction. Nutrient agar plus 2 per cent glucose: Colonies appear in 36 hours. After 3 days colonies circular, smooth, glistening, convex; edges entire; light pink, but not constant. Broth : Good growth. Pellicle and sediment. Milk : Little change, if an.y. Nitrites not produced from nitrates. Indole not formed. No HoS produced. Acid from arabinose, glucose, galac- tose, fructose, sucrose and glycerol. No acid from lactose, maltose, dextrin and inulin. Starch not hydrolyzed. Optimum temperature 21° to 25°C. Source : Description based on 7 cul- tures isolated from rotting apples and from apple maggots. Habitat : Pathogenic on apples, and found with the apple maggot, Rhagoletis pomonella. 139. Pseudomonas helianthi (Kawa- mura) comb. nov. (Bacterium helianthi Kawamura, Ann. Phyt. Soc. Japan, 4, 1934, 27; Phytomonas helianthi Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 362.) From M. L. Helian- thus, a generic name. Probable synonym : Phytomonas helian- thi var. tubero.si Thornberry and Ander- son, Phytopath., 27, 1937, 948. Rods: 1 to 1.4 by 1.6 to 2.4 microns. Motile with a single polar flagellum. Gram-negative. Gelatin: No liquefaction. Beef agar colonies: White, circular, edges entire. Broth: Turbid. Pellicle. Milk: Peptonized. Litmus reduced. Nitrates: Gas production. Indole not produced. No HoS produced. Acid but not gas from sucrose and glyc- erol. No acid from lactose and maltose. Starch hydrolyzed. Optimum temperature 27° to 28°C. Maximum 35.5°C. Minimum 12°C. Good growth at pH 6.4. No growth pH 5.4 and pH 8.8. Habitat : Pathogenic on sunflower, Helianthus debilis. 140. Pseudomonas alboprecipitans Rosen. (Rosen, Ann. Missouri Bot. 142 MANUAL OF DETERMINATIVE BACTERIOLOGY Garden, 9, 1922, 383; Bacterium albo- precipitans Elliott, Bact. Plant Path., 1930, 89; Phytomonas alboprecipitans Bergey et al., Manual, 3rd ed., 1930, 277.) From Latin albus, white and precipitans , precipitating, referring to the white precipitate produced in cul- ture. Rods: 0.6 by 1.8 microns, occurring singly or in pairs. Capsules present. Motile with a polar flagellum. Gram- negative. Gelatin: Not liquefied. Nutrient agar colonies : White, circular, raised, smooth, sticky, with margins entire. Whitish discoloration of the medium. Broth: Turbid in 24 hours. Heavy sediment in old cultures. Milk : Becomes alkaline and slowly clears. Nitrites produced from nitrates. Indole not produced. No HzS produced. Acid but not gas from glucose, fruc- tose, glycerol and mannitol. No acid from lactose, maltose or sucrose. Starch is hydrolyzed. Optimum temperature 30° to 35°C. Maximum temperature 40 °C. Minimum 0°C. Aerobic. Distinctive characteristics : White pre- cipitate in culture media. Source : Isolated a number of times from foxtail grass. Habitat : Pathogenic on foxtail, Chaeto- chloa lutescens and other grasses. 141. Pseudomonas petasitis (Taki- moto) comb. nov. {Bacterium petasitis Takimoto, Ann. Phyt. Soc. Japan, 2, 1927, 55; Phytomonas petasitis Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 393.) From M. L. Petasites, a generic name. Rods: 0.8 to 1.1 by 1.1 to 1.7 microns. Motile with a polar flagellum. Gram- negative. Gelatin: No liquefaction. Beef agar colonies : White, circular or ameboid, butyrous. Broth: Strong turbidity. Pellicle. Milk: Coagulated in 30 days. Nitrites produced from nitrates with gas formation. Indole not produced. No HoS produced. No evident acid in peptone broth but gas from glucose, lactose and sucrose. .\cid but not gas from glycerol. Weak growth in broth plus 6 percent salt. Optimum temperature 27° to 30 °C. Maximum 47°C. Minimum approxi- mately 5°C. Source: Isolated from brown to black lesions on Petasites japonicus in Japan. Habitat : Pathogenic on leaves of Petasites japonicus. 142. Pseudomonas lignicola Wester- dijk and Buisman. (De lepenziekte, Arnliem, 1929, 51.) From Latin, lignum, wood; -cola, dweller. Rods: Single or short chains. Motile with 1 to several polar flagella. Gram- negative. Gelatin: No liquefaction. Malt agar streaks : Milk white with a colorless edge. Broth : Turbid with light pellicle. Milk: No coagulation. No acid. Nitrites not produced from nitrates. Indole not formed. Starch hydrolysis slight. Optimum temperature ±25°C. Source : From vessels of elm wood showing dark discoloration, in Holland. Habitat: Pathogenic in elm wood. 143. Pseudomonas andropogoni (Erw. Smith) Stapp. {Bacterium andropo- goni Erw. Smith, Bacteria in Relation to Plant Diseases, 2, 1911, 63; Elliott and Smith, Jour. Agr. Res., 38, 1929, 4; Stapp, in Sorauer, Handbuch der Pflanzen- krankheiten, 2, 5 Auf., 1928, 27; Phyto- monas andropogoni Bergey et al., Manual, 3rd ed., 1930, 276.) From M. FAMILY PSEUDOMONADACEAE 143 L. Andropogon , a generic name (a syn- onym of Holcvs). Description from Elliott and Smith (loc. cit.). Rods: 0.64 by 1.76 microns. Motile with one to several bipolar flagella. Capsules. Gram-negative. Gelatin: Feeble liquefaction or none. Beef -extract agar colonies : Slow -grow- ing, round, smooth, glistening, viscid, white. Broth : Growth slow with moderate turbidity in 48 hours. A thin pellicle. Milk: Alkaline and clears. Nitrites not produced from nitrates. Indole not formed. No H2S formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from glucose, arabi- nose, fructose and xylose. No acid from .sucrose, maltose, lactose, raffinose, glyc- erol and mannitol. Starch partially digested. Optimum temperature 22' to 30°C. Ma.ximum 37° to 38 °C. Minimum 1.5°C. Optimum pH 6.0 to 6.6. Maximum 8.3 to 8.6. Minimum 5.0. Source : Elliott used for her description 4 cultures isolated from lesions on sorgo, sorghum and broom-corn. Habitat : Pathogenic on sorghum, Hol- cus sorghum. 144. Pseudomonas woodsii (Smith) Stevens. {Bacterium woodsii Erw. Smith, Bact. in Relation to Plant Dis- eases, 2, 1911, 62; Stevens, Plant Disease Fungi, New York, 1925, 39; Phytomonas woodsii Bergey et al., Manual, 3rd ed., 1930, 256.) Named for A. F. Woods, American plant pathologist. Description from Burkholder and Gu- terman, Phytopath., 25, 1935, 118. Rods: 0.67 by 1.56 microns. Motile with a polar flagellum. Gram-negative. Gelatin: No liquefaction. Beef -extract agar slants : Growth slow and scant, filiform, creamy, butyrous. Broth: Turbid. Milk: Becomes alkaline but otherwise little changed. Nitrites not produced from nitrates. Indole not formed. No HoS formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 601). Acid but not gas from glucose, fruc- tose, galactose, arabinose, xylose, rham- nose, lactose, glycerol and mannitol. Alkaline reaction from salts of acetic, citric, malic and succinic acids. Sucrose, maltose, salicin, and lactic and formic acids not fermented. Starch not hy- drolyzed. Slight growth in broth plus 3 per cent salt. Aerobic. Source : Isolated from water-soaked lesions on carnation leaves. Habitat: Pathogenic on carnation, Dianthus caryophyllvs. 145. Pseudomonas panici-miliacei (Ikata and Yamauchi) comb. nov. {Bac- terium panici-jriiliacei Ikata and Yamau- chi, Jour. Plant Protect., 18, 1931, 35; Phytomonas panici-miliacei Burkholder, in Manual, 5th ed., 1939, 204.) From M. L. Panicum miliaceum. Description translated by Dr. K. Togas hi. Rods: 0.8 to 1.1 by 1.8 to 2.6 microns. Motile, with a single flagellum. Gram- negative. Gelatin : Not liquefied. Potato-agar plates : Growth moderate, whitish, then tinged with light orange, undulating margins. Broth: Turbid, white pellicle formed. Milk: No coagulation and slow diges- tion. Alkaline. Nitrites are produced from nitrates. Indole not formed. No H2S produced. No acid and no gas from sucrose, glu- cose, lactose, glycerol and sodium nitrate. Starch not hydrolyzed. Optimum temperature 30° to SS'C. 144 MANUAL OF DETERMINATIVE BACTERIOLOGY Facultative anaerobe. Source : Species first isolated from millet, Panicum miliaceum. Habitat : Causes a leaf stripe of Pani- cum miliaceum. 146. Pseudomonas saliciperda Lindei- jer. (Lindeijer, Inaug. Diss., Univ. Amsterdam, 1932; Phytopath. Ztschr., 6, 1933, 373; Bacterium saliciperda Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 106; Phytomonas salici- perda Magrou, in Hauduroy et al., Diet, d. Bact. Path., Paris, 1937, 408.) From L. salix (salicis), willow; perdo, to destroy. Rods: 1.2 to 2.1 microns in length. Motile with a polar fiagellum. Gram- negative. Gelatin: No liquefaction. Beef wort agar colonies: Gray -white. Milk: No acid nor coagulation. Nitrites produced (small amount) from nitrates. Indole formation slight. No gas from carbohydrates. Starch not hydrolyzed. Facultative anaerobe. Source: Isolated from wilted branches of willow and pathogenicity proved. Habitat : Pathogenic on willow, Salix spp. 147. Pseudomonas eriobotryae (Taki- moto) Dowson. (Bacterium eriobotryae Takimoto, Jour. Plant Protect., 18, 1931, 354; Phytomonas eriobotryae Burkholder, in Manual, 5th ed., 1939, 205; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 10.) From M. L. Eriobotrya, a generic name. Translated by Dr. K. Togashi. Rods: 0.7 to 0.9 by 2.2 to 3.0 microns. Motile, w^ith 1 or 2 flagella. Gram- negative. Gelatin: Not liquefied. Agar-plates : Colonies appear after 3 days, white or hyaline, butyrous, margins entire. Broth: Moderately turbid, pellicle powdery, ring formed. Milk : No coagulation, peptonized slowly. Alkaline. Nitrites not produced from nitrates. Indole not formed. No H^S produced. No acid or gas from glucose, sucrose, lactose and glycerol in broth. Starch not hydrolyzed. Temperature relations : Minimum below 4°C, optimum 25° to 26°C, and maximum 32°C. Thermal death point 51°C. Aerobic. Source : Species isolated from loquat, Eriobotrya japonica. Habitat : Causes a bud rot of Eriobotrya japonica. 148. Pseudomonas wieringae (Elliott) comb. nov. {Phytomonas betac Wieringa, Nederl. Tijdschr. Hyg., Microbiol, en Serol., Leiden, 2, 1927, 148; Bacterium wieringae Elliott, Man. Bact. Plant Pathogens, 1930, 264; Phytomonas wier- ingae Burkholder, in Manual, 5th ed., 1939, 206.) Named for K. L. Wieringa, plant pathologist of Holland. Because Bacterium betae Chester (Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 53) may be a pseudomonad, the species name proposed by Elliott has been retained. Description from Elliott (loc. cit.). Rods : 0.5 to 2.0 microns. Motile with 1 to 5 polar flagella. Gram-negative. Beef-agar colonies: Smooth, round, white to grayish, fluorescent. Milk: Cleared in 5 days. Not co- agulated. Nitrites not produced from nitrates. No gas from sugars. Optimum temperature 28° to 30 °C. Maximum 37°C. Minimum 4°C. Source : Isolated from vascular rot of beets in Holland. Habitat : Pathogenic on beets, Beta vulgaris. FAMILY PSEUDOMONADACEAE 145 Appendix I*: The following species are believed to belong in the genus Pseudo- monas although descriptions are fre- (luently incomplete. Achromohacter pellucidinn Harrison. (Canadian Jour. Res., 1, 1929, 236.) Isolated from halibut. For a description of this species, see Bergey et al., Manual, othed., 1939, 619. Bacillus aurantiacus tingitanus Rem- linger and Bailly. (jNIaroc Medical, No. 150, 1935; See Lasseur, Dupaix and Babou, Trav. Lab. Microbiol. Fac. Pharm. Nancy, Fasc. 8, 1935, 41.) From water. Dissociates readily. Related to Pseudomonas fluorescens aureus Zim- mermann. See p. 645. Bacillus cyaneofluorescens Zaugemeis- ter. (Cent, f . Bakt., I Abt., 18, 1895, 321 ; Pseudomonas cyaneofluorescens Migula, Syst. d. Bakt., 2, 1900, 906.) From blue milk. Bacillus fluorescens nivalis Eisenberg. (Eine Gletscherbakterie, Schmelck, Cent. f. Bakt., 4, 1888, 545; Eisenberg, Bakt. Diag., 3 Aufl., 1891, 77.) From the melting snow of a glacier. Probably a S3'nonym of Pseudomonas fluorescens. Bacillus lactis saponacei Weigmanii and Zirn. (Cent. f. Bakt., 15, 1894, 468. * From soapy milk. Bacterium auxinophilum Jacobs. (Ann. Appl. Biol., 23, 1935, 619.) A Gram-negative organism with a polar flagellum which liquefies gelatin rapidly. Bacterium bosporum Kalnins. (Lat- vijas Universitates Raksti, Serija I, No. 11, 1930, 259.) Decomposes cellu- lose. Single polar flagellum. From soil. Bacterium briosii FsLvavino. (Atti 1st. Bot. d. R. Univ. di Pavia, Ser. 2, 12, 1910, .337.) The natural host is Lyco- persicon esculentum. Pavarino (Rev. di Patol. Veg., 6, 1913, 161) states that this organism and Phytobacter lycopersicuni Groenewege (Meded. Rijks. Hoogere Land, Tuin- en Boschbouwschool, Dell 5, 5, 1912, 217) should be considered identi- cal. It closely resembles Bacterium vesicatoriuin Doidge (Jour. Dept. Agr. So. Africa, 1, 1920, 718) according to Gardner and Kendrick (Jour. Agr. Res., 21, 1921, 140). Bacterium daphorum Kalnins. (Lat- vijas Universitates Raksti, Serija I, No. 11, 1930, 257.) Decomposes cellu- lose. Single polar flagellum. From soil. Bacterium fraenkelii Hashimoto. (Zeit. f. Hyg., 31, 1899, 88.) A pleomorphic polar flagellate bacterium. From milk. Bacterium gummis Comes. (Comes, Napoli, Maggio 18, 1884, 14; see Comes, Atti d. R. 1st. d'incoraggiamento alii Sc, Ser. 3, 3, 1884, 4; Bacillus gummis Trevisan, I generi e le specie delle Batteriacee, Milano, 1889, 17.) Patho- genic on grapes, Viiis spp. Bacterium krameriani Pavarino. (Atti R. Accad. Naz. Lincei Rend. CI. Sci. Fis., Mat. et Nat., 20, 1911, 233.) Pathogenic on the orchid, Oncidium krameriani. Bacterium pusiolum Kalnins. (Lat- vijas U'niversitates Raksti, Serija I, No. 11, 1930, 261.) Decomposes cellu- lose. Single polar flagellum. From manure. Bacterium protozoides Kalnins. (Lat- \'ijas U'niversitates Raksti, Serija I, No. 11, 1930, 263.) Decomposes cellu- lose. Single polar flagellum. From soil. Pseudomonas acuta INIigula. (Culture No. 11, Lembke, Arch. f. Hyg., 29, 1897, 317 ; Migula, Syst. d. Bakt., ~2, 1900, 921 .) From the intestine. Pseudomonas alba Migula. {Bacillus fluorescens albus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, I Reihe, 1890, 18; Migula, Syst. d. Bakt., 2, 1900, 909.) From water. Bacillus fluorescens non liquefaciens Eisenberg, Bakt. Diag., 3 Aufl., 1891, 145 may be identical ac- cording to Migula {loc. cit.). Pseudomonas allii (Griffiths) Migula. {Bacterium allium Griffiths, Proc. Roy. Soc. Edinburgh, 51, 1887, 40; Migula. * Appendixes I and II prepared by Prof. Robert S. Breed, New York State Experi- ment Station, Geneva, New York, July, 1943. 146 MANUAL OF DETERMINATIVE BACTERIOLOGY Syst. d. Bakt., 2, 1900, 932.) From rotted onions. Pseudomonas aquatilis Migula. (Tata- roff, Inaug. Diss., Dorpat, 1891, 31; Migula, Syst. d. Bakt., 2, 1900, 933.) From water. Said to form spores. Pseudomonas aromatica Migula. (Bacillus crassus aromaticus Tataroff, Inaug. Diss., Dorpat, 1891, 27; Migula, Syst. d. Bakt., 2, 1900, 880.) From water. Pseudomonas aromatica var. quercilo- pyrogallica Kluyver, Hof and Boe- zaardt. (Enzymologia, 7 1939, 28.) Pseudomonas arliculaia Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 462.) From the stomachs of birds. Pseudomonas aucubicola Trapp. (Phytopath., 26, 1936, 264.) Isolated from Aucuba japonica. Not pathogenic. Pseudomonas aurea Migula. (Bacillus fluorescens aureus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, I Reihe, 1890, 14; Migula, Syst. d. Bakt., 2, 1900, 931.) From water. Pseudomonas brassicae acidae G ruber. (Cent. f. Bakt., II Abt., 22, 1909, 558). From sauerkraut. Identical with Bac- terium brassicae acidae Conrad (Arch. f. Hyg., 29, 1897, 75) according to Lehmann and Neumann (Bakt. Diag., 5 Aufi., 2 1912, 380.) Pseudomonas bulyri Migula. (Bacil- lus bulyri fluorescens Lafar, Arcli. f. Hyg., 13, 1891, 19; Migula, Syst. d. Bakt., 2, 1900, 894.) From butter. Pseudomonas calciphila Molisch. (Cent. f. Bakt., II Abt., 65, 1925, 136.) From fresh water. Deposits CaCOa. Pseudomonas calco-acetica Clifton. (Enzymologia, 4, 1937, 246.) Pseudomonas capsulata Migula. (Syst. d. Bakt., 2, 1900, 915; not Pseudo- monas capsulata Bergey et al., Manual, 1st ed., 1923, 124.) Similar to Pseudo- monas macroselmis Migula. Pseudomonas caryocyanea (Dupaix) Beijerinck. (Bacillus caryocyaneus Du- paix, Thesis, Univ. of Nancy, 1933, 1 ; Beijerinck, see Dupaix', ibid., 13; Bac- terium caryocyaneum Dupaix, ibid., 246.) Isolated from rotten willow wood, from yeast mash and beer-wort. Name ap- pears first as Bacillus caryocyaneus on a culture sent by Beijerinck from Delft, Holland to the National Collection of Type Cultures, Lister Institute, London. Regarded by Dupaix as closely related to Bacillus cyaneo -fluorescens Zangemeister (Cent. f. Bakt., I Abt., 18, 1895, 321; Pseudomonas cyaneo -fluorescens Migula, Syst. d. Bakt., 2, 1900, 906); Der blaue Bacillus, Mildenberg (Cent. f. Bakt., II Abt., 56, 1922, 309; Pseudomonas mildenbergii Bergey et al., Manual, 3rd ed., 1930, 172); and Bacillus pyocyaneus Gessard (Compt. rend. Acad. Sci. Paris, 94, 1882,536). Pseudovionas catarrhalis Chester. (Bacillus der Hundestaupe, Jess, Cent, f . Bakt., II Abt., 25, 1899, 541; Chester, Man. Determ. Bact., 1901 , 308.) Isolated from catarrh in dogs. Pseudomonas caviae Scherago. (Jour. Bact., 31, 1936, 83; Jour. Inf. Dis., 60, 1937, 245.) Cause of an epizootic septi- cemia in guinea pigs. Pseudomonas chlorophaena Migula. (Syst. d. Bakt.,^, 1900, 899.) Pseudomonas coccacea Migula. (Cul- ture No. 10, Lembke, Arch. f. Hyg., 29, 1897, 317; Migula, Syst. d. Bakt., 2, 1900,924.) From the intestine. Pseudomonas cohaerea (sic) (Wright) Chester. (Bacillus cohaerens Wright, Mem. Nat. Acad. Sci., 7, 1895, 464; Chester, Man. Determ. Bact., 1901, 312.) From water. Pseudomonas coli Migula. (Culture No. 8, Lembke, Arch. f. Hyg., 29, 1897, 315; Migula, Syst. d. Bakt., 2, 1900, 920.) From the intestine. Pseudomonas colloides Migula. (Bacillus fluorescens putidus colloides Tataroff, Inaug. Diss., Dorpat, 1891, 40; Migula, Syst. d. Bakt., 2, 1900, 902.) From water. Said to form spores. Pseudomonas conradi Lehmann and Neumann. (Bakt. Diag., 5 Aufl., 2, 1912, 394.) Red pigment. FAMILY PSEUDOMONADACEAE u: Pseudonionas delabcns (Wright J Chester. (Bacillus delabens Wright, Mem. Nat. Acad. Sci., 7, 1895, 456; Chester, Man. Determ. Bact., 1901, 314.) From water. Pseudonionas duplex Migula. (Cul- ture No. 7, Lembke, Arch. f. Hyg., 29, 1897, 314; Migula, Syst. d. Bakt., 2, 1900,922.) From the intestine. Pseudotnonas cllipsoiden Migula. (Bacillus oogenes fluorcsccns 0, Zorken- dorfer, Arch. f. Hyg., 16, 1893, 393; Migula, Syst. d. Bakt., 2, 1900, 925.) From hens' eggs. Pseudomonas ephemerocyanea Fuller and Norman. (Jour. Bact., Ji.6, 1943, 274.) From soil. Decomposes cel- lulose. Pseudomonas erylhra Fuller and Nor- man. (Jour. Bact., 46, 1943, 276.) From soil. Decomposes cellulose. Pseudomonas erythrospora (Cohn) Migula. (Bacillus erylhrosporus Cohn, Beitr. z. Biol. d. Pfianzen, S, Heft 1, 1879, 128; Migula, in Engler and Prantl, Die nautrl. Pflanzenfam., 1, la, 1895, 29.) From air, meat infusion and water. Said to form spores. Pseudomonas fimhriata (Wright) Chester. (Bacillus fimbriatus Wright, Mem. Nat. Acad. Sci., 7, 1895, 463; Chester, Man. Determ. Bact., 1901, 313.) From water. Pseudomonas fluorcsccns cxiliosus van Hall. (Ztschr. f. Pflanzenkr., IS, 1903, 132.) Causes soft rot of shoots and bulbs of iris (Iris spp.). Pseudomonas foliacea Chester. (Bacillus fluorcsccns foliaceus Wright, Mem. Nat. Acad. Sci., 7, 1895, 439; Chester, Man. Determ. Bact., 1901, 324; Bacillus fluorescens-foliaceus Chester, ibid.) From water. Very similar to Pseudomonas incognita Chester. Pseudomonas gasoformans ]\Iigula. (Ein neuer gasbildender Bacillus, Gart- ner, Cent. f. Bakt., 15, 1894, 1; Migula, Syst. d. Bakt., 2, 1900, 883.) Gas bubbles in gelatin stab. Pseudomonas gracilis Migula. (Syst. d. Bakt., 8, 1900, 888.) Morphologically like Pseudomonas fluorescens Migula. Pseudomonas granulata Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 464.) From the stomach and intestine of birds. Pseudomonas halestorgus Elazari- Volcani. (Ph.D. Thesis, Hebrew Univ., Jerusalem, 1940.) A halophilic pseudo- monad from the Dead Sea. Pseiido7no7ias hydrosulfurea Migula. (Bacillus oogenes hydrosidfureus fi, Zor- kendorfer, Arch. f. Hyg., 16, 1893, 385; Migula, Syst. d. Bakt., 2, 1900, 898.) From hens' eggs. Pseudomonas iridis van Hall. (Van Hall, Thesis, Univ. Amsterdam, 1902 and Ztschr. f. Pflanzenkr., 13, 1903, 129; Bacterium iridis Elliott, Man. Bact. Plant Path., 1930, 142; Phytomonas iridis Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 369.) Causes a rot of bulbs and leaves of iris (7ns spp.). Pseudoinonas iris (Frick) Migula. (Bacillus iris Frick, Arch. f. path. Anat., 116, 1889, 292; according to Eisenberg, Bakt. Diag., 3 Aufl., 1891, 148; Migula, Syst. d. Bakt., ^,1900,931.) Pseudomonas ilalica (Foa and Chia- pella) Reinelt. (Quoted from Lehmann and Neumann, Bakt. Diag., 7 Aufi., 2, 1927, 367.) Phosphorescent. Pseudomonas javanica (Eijkmann) Migula. (Photobacterium javanense Eijkmann, Geneesk. Tijdschr. v. Nederl. -Indie, 32, 1892, 109; Abst. in Cent. f. Bakt., 12, 1892, 656; Bacillus java7iiensis Dyar, Annals New York Acad. Sci., 8, 1895, 359; Bacterium java- niensis Chester, Man. Determ. Bact., 1901, 170; Photobacterium javanicum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 199; Migula, Syst. d. Bakt., 2, 1900, 953.) From sea fish in Java. Blue-green luminescence. Pseudomonas lactica Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 238.) From a vegetable infusion. Pseudomonas lasia Fuller and Norman. 148 MANUAL OF DETERMINATIVE BACTERIOLOGY fJour. Bact., 46, 1943, 275.) From soil. Decomposes cellulose. Pseudomonas lembkei Migula. (Cul- ture No. 12, Lembke, Arch. f. Hyg., 29, 1897, 318; Migula, Syst. d. Bakt., 2, 1900, 896.) From the intestine. Pseudomonas liquefaciens (Tataroff) Migula. {Bacillus liquefaciens Tataroff, Inaug. Diss., Dorpat, 1891, 29; Migula, Syst. d. Bakt., 2, 1900, 876.) From water. Pseudomonas listeri Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 260.) From a vegetable infusion. Pseudomonas Zong^a Migula. {Bacillus fluorescens longus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, I Reihe, 1890, 20; Migula, Syst. d. Bakt., 2, 1900, 907.) From water. Pseudomonas macruselmis Migula. {Bacillus Jlvorescens putidus Tataroff, Inaug. Diss., Dorpat, 1891, 42; Migula, in Engler and Prantl, Die natlirl. Pflan- zenfam., 1, la, 1895, 29.) From water. Pseudomonas maidis (Eisenberg) Migula. {Bacillus maidis Eisenberg, Bakt. Diag., 3 Aufl., 1891, 119; Migula, Syst. d. Bakt., 2, 1900, 877.) From corn grains soaked in water and from feces of pellagra patients. Pseudomonas maschekii Migula. (Blaugriiner Bacillus, Maschek, Bakt. Untersuch. d. Leitmeritzer Trinkwasser, Jahresber. d. Oberrealschule zu Leit- meritz, 1887; Migula, Syst. d. Bakt. 2, 1900, 916.) From water. Pseudomonas melochlora (Winkler and Schrotter) Migula. {Bacillus melo- chloros Winkler and Schrotter, Ein neuer grunen Farbstoff entwickelnder Bacillus, Wien, 1890; Migula, Syst. d. Bakt., 2, 1900, 893.) From caterpillar feces. Pseudomonas mesenterica Migula. {Bacillus fluorescens mesentericus Tata- roff, Inaug. Diss., Dorpat, 1891, 38; Migula, Syst. d. Bakt., 2, 1900, 903.) From water. Pseudomonas metalloides Migula. {Bacillus rosaceus metalloides Tataroff, Inaug. Diss., Dorpat, 1891, 65; not Bac- terium rosaceus metalloides Dowdeswell, Ann. de Microgr., 1, 1888-89, 310, see Heffernan, Cent. f. Bakt., II Abt., 8, 1902, 689; Pseudomonas rosacea Migula, in Engler and Prantl, Die natlirl. Pflan- zenfam., 1, la, 1895, 29; Migula, Syst. d. Bakt., 2, 1900, 938.) Single flagellum. Red and yellow -red pigment. From water. Pseudomonas minutissima Migula. {Bacillus fluorescens liquefaciens minutissimus Unna and Tommasoli, Monatsh. f. prakt. Dermat., 8, 1889, 57; according to Eisenberg, Bakt. Diag., 3 Aufl., 1891, 76; Migula, Syst. d. Bakt., 2, 1900, 891.) Found on human skin in cases of seborrheic eczema. Pseudomonas mohilis Migula. (Cul- ture No. 9, Lembke, Arch. f. Hyg., 29, 1897, 316; Migula, Syst. d. Bakt., 2, 1900,923.) From the intestine. Pseudomonas monadiformis (Kruse) Chester. {Bacillus coli mohilis Messea, Riv. d'Igiene, Rome, 1890; Bacillus monadiformis Kruse, in Fltigge, Die Mikroorganismen, 2, 1896, 374; Chester, Man. Determ. Bact., 1901, 308.) From typhoid stools. Pseudomonas mucidolens Levine and Anderson. (Jour. Bact., 2S, 1932, 337.) Causes musty odors in eggs. Also milk (Olsen and Hammer, Iowa State Coll. Jour. Sci., 5, 1934,125). Pseudomonas mucidolens var. tarda Levine and Anderson. (Jour. Bact., 23, 1932, 337.) Causes mustj'- odors in eggs. Pseudomonas nexibilis (Wright) Chester. {Bacillus nexibilis Wright, Mem. Nat. Acad. Sci., 7, 1895, 441; Chester, Man. Determ. Bact., 1901, 309.) From water. Pseudomonas nivalis Szilvinyi. (Cent. f. Bakt., II- Abt., 94, 1936, 216.) A red chromogen isolated from red snow in Austria. Pseudomonas ochroleuca Migula. {Bacillus y, Zorkendorfer, Arch. f. Hyg., 16, 1893, 396; Migula, Syst. d. Bakt., 2, 1900, 897.) From hens' eggs. Pseudomonas oogenes Migula. {Bacil- lus oogenes hydrosidfureus 8, Zorken- dorfer, Arch. f. Hyg., 16, 1893, 386; FAMILY PSEUDOMONADACEAE 149 Migula, Syst. d. Bakt., 2, 1900, 878.) Single flagellum. From hens' eggs. Pseudomonas ovi Migula. (Bacillia^ oogenes fluorescens e, Zorkendorfer, Arch, f. Hyg., 16, 1893, 395; Migula, Syst. d. Bakt., 2, 1900, 924.) From hens' eggs. Pseudomonas ovicola Migula. {Bacil- lus oogenes fluorescens y, Zorkendorfer, Arch. f. Hyg., 16, 1893, 394; Migula, Syst. d. Bact., 2, 1900, 925.) From hens' Pseudomonas pallescens Migula. (Bacillus viridis pallescens Frick, in Virchow, Arch. f. path. Anat., 116, 1889, 292; according to Eisenberg, Bakt. Diag., 3 Aufl., 1891, 154; Migula, Syst. d. Bakt., 2, 1900, 927.) Source not given. Pseudomonas pansinii Migula. (Bacillus fluorescens non liquefaciens Pansini, in Vixchow, Arch. f. path. Anat., 122, 1890, 452; Migula, Syst.-d. Bakt., 2, 1900, 926.) Pseudomonas pelliculosa Migula. (Bacillus oogenes fluorescens d, Zorken- dorfer, Arch. f. Hyg., 16, 1893, 395; Migula, Syst. d. Bakt., 2, 1900, 926.) From hens' eggs. Pseudomonas pellucida Kern. (.Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1896, 465.) From the intestine of birds. Pseudomonas plehniae Spieckermann and Thienemann. (Arch. f. Hyg., 74, 1911, 110.) Isolated from carp. Patho- genic for many species of fish. Pseudomonas plicala (Frankland and Frankland) Migula. (Bacillus plicatus Frankland and Frankland, Phil. Trans. Roy Soc. London, 178, B, 1887, 273; Migula, Syst. d. Bakt., 2, 1900, 881.) From air. Pseudomonas pseudolyphosa Migula. (Typhusahnlicher Bacillus, Lustig, Diag. d. Bakt. d. Wassers, 1893, 16; Migula, Syst. d. Bakt., 2, 1900, 893.) From water. Pseudomonas pidlulans (Wright) Chester. (Bacillus pullidans Wright, Mem. Nat. Acad. Sci., 7, 1894, 445; Chester, Man. Determ. Bact., 1901, 315.) From water. Pseudomonas prolea Frost. (U. S. PubHc Health Ser., Hyg. Lab. Bull. 66, 1910, 27.) From filtered river water. Can be agglutinated by specific typhoid immune-serum. Pseudomonas rosea Chester. (Bacil- lus roseus vini Bordas, Joulin and Rackowski, Compt. rend. Acad. Sci. Paris, 126, 1898, 1550; Chester, Man. Determ. Bact., 1901, 327; not Pseudo- monas rosea Migula, in Engler and Prantl, Dienaturl.Pfllanzenfam.,i, la, 1895,30.) From wine. Said to form spores. Pseudomonas (Hydrogenomonas) sac- charophila Doudoroff. (Enzymologia, 9, 1940, 50.) From stagnant water. Pseudomonas sapolactica (Eichholz) De Rossi. (Bacterium sapolacticum Eichholz, Cent. f. Bakt., II Abt., 9, 1902, 631; De Rossi, Microbiologia Agraria e Technica, Torino, 1927, 693.) Isolated from soapy milk. Pseudomonas sencea Migula. (Seiden- glanzender Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 26; Migulu, Syst. d. Bakt., 2, 1900, 882.) Pseudomonas tenuis Migula. (Bacil- lus fluorescens tenuis Zimmermann, Bakt. unserer Trink- u. Nutzwasser, I Reihe, 1890, 16; Migula, Syst. d. Bakt., 2. 1900,910.) From water. Pseudomonas Irommehchldgel (Rav- enel) Chester. (Bacillus iromnielschldgel Ravenel, INIem. Nat. Acad. Sci., 8, 1896, 26; Chester, Man. Determ. Bact., 1901, 327.) From soil. Said to form spores. Pseudomonas undidala Chester. (Bacillus fluorescens unditlalus Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 20; Chester, Man. Determ. Bact., 1901, 328.) From soil. Said to form spores. Pseudomonas virescens (Frick) Migula. (Bacillus virescens Frick, Arch. f. path. Anat., 116, 1889, 292; Migula, Syst. d. Bakt., ^,1900,916.) From green sputum. Pseudomonas viridans (Symmers) Migula. (Bacillus viridans Symmers, Brit. Med. Jour., No. 1615, 1891, 1252; Abst. in Cent. f. Bakt., 12, 1892, 165; Migula, Syst. d. Bakt., 2, 1900, 890.) From water. 150 MANUAL OF DETERMINATIVE BACTERIOLOGY Psctidorunnas viridescens Chester. (Bacillus viridescens liquefaciens Rav- enel, Mem. Nat. Acad. Sci., 8, 1896, 24; Chester, Man. Determ. Bact., 1901, 328.) From soil. Said to form spores. Pseudomonas viridis Migula. (Bacil- lis der grimen Diarrhoe der Kinder, Lesage, Arch. d. Physiol, norm, et path., 20, 1888, 212; see Eisenberg, Bakt. Diag., 3 Aufl., 1891. 238; Miguhi, Syst. d. Bakt., 2, 1900, 886.) From intestine of children. Pseudomonas weigmanni Migula. (Bakterie IV, Weigmann and Zirn, Cent, f. Bakt., 15, 1894, 466; Migula, Syst. d. Bakt., 2, 1900, 892.) From soapy milk. Pseudomonas zorkendorferi Migula. (Bacillus oogenes fluoresccns a, Zorken- dorfer. Arch. f. Hyg., 16, 1893, 392; Migula, Syst. d. Bakt., 2, 1900, 897.) From hens' eggs. Appendix II : The following polar flagel- late organism has been described from activated sludge. H. Winogradsky has also described polar flagellate forms from the same source that form zoogloea (Compt. rend. Acad. Sci. Paris, 200, 1935, 1887; Ann. Inst. Pasteur, 58, 1937, 333). Zoogloea ramigera Kruse emend. Butterfield. (Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 1, 1896, 68; Butterfield, Public Health Reports, 50, 1935, 671; Culture No. 50, Wattle, Pub. Health Reports, 57, 1942, 1519.) Rods: 1 by 2 to 4 microns, with rounded ends. Non-spore-forming. Capsules present. Motile with a single long polar flagellum. Gram-negative. Gelatin : No liquefaction. Grows better in aerated liquid media. Agar : Scant growth. Indole not formed. No HoS produced. No acid or gas from carbohydrates. Nitrites not produced from nitrates. Optimum pH 7.0 to 7.4. Optimum temperature 28° to 30 °C. Good growth at 20° and at 37 °C. Mini- mum temperature 4°C. Strict aerobe. Distinctive character : Oxidizes sewage. Source : Isolated from activated sludge. Habitat: Produces zoogloeal masses in activated sludge. Genus II. Xanthomonas Dowson* (Phytoinonas Bergey et al., Manual, 1st ed., 1923, 174 ; Dowson, Cent, f . Bakt., II Abt., 100, 1939, 187.) Cells usually monotrichous, with yellow, water-insoluble pigment. Proteins are usually readily digested. Milk usually becomes alkaline. Hydrogen sulfide is pro- duced. Asparagin is not sufficient as an only source of carbon and nitrogen. Acid is produced from mono- and disaccharides. Mostly plant pathogens causing necrosis. From Gr. xanthus, yellow ; monas, a unit ; M. L. monad. The type species is Xanthomonas hyacinihi (Wakker) Dowson. Key to the species of genus Xanthomonas. 1. Colonies yellow. a. Gelatin liquefied. b. Starch hydrolysis feeble. c. Nitrites not produced from nitrates. 1. Xanthomonas hyacinthi. 2. Xanthomonas pruni. 3. Xanthomonas vitians. Prepared by Prof. Walter H. Burkhol der, Cornell Univ., Ithaca, N. Y., June, 1943. FAMILY PSEUDOMONADACEAE 151 cc. Nitrites produced from nitrates. 4. Xanthomonas beticola. 5. Xanthomonas lactucae-scariolac. 6. Xanthomonas ruhrilincans. bb. Starch hydrolysis strong. c. Nitrites not produced from nitrates. d. No brown pigment in beef-extract agar. 7. Xanthomonas barbareae. 8. Xanthomonas begoniac. 9. Xanthomonas campestris. 9a. Xanthomonas campestris var. armoraciae. 10. Xanthomonas citri. 11. Xanthomonas corylina. 12. Xanthomonas cucurbitae. 13. Xanthomonas dieffenbachiae. 14. Xanthomonas holcicola. 15. Xanthomonas incanae. 16. Xanthomonas jitglandis. 17. Xanthomonas lespedezae. 18. Xanthomonas malvacearnm. 19. Xanthomonas pelargonii. 20. Xanthomonas phaseoli. 20a. Xanthomonas phaseoli var. sojense. 21. Xanthomonas plantaginis. 22. Xanthomonas ricinicola. 23a. Xanthomonas translucens f. sp. hordei. 23b. Xanthomonas translucens i. sp. undidosa. 23c. Xanthomonas translucens f. sp. secalis. 23d. Xanthomonas translucens f. sp. hordei-avenae. 23e. Xahthomonas translucens f. sp. cerealis. 24. Xanthomonas vascular inn. 25. Xanthomonas vesicatoria. 25a. Xanthomonas vesicatoria var. raphani. dd. Brown pigment produced in beef -extract media. 26. Xanthomonas nakalac. 20b. Xanthomonas phaseoli var. fuscans. cc. Nitrites produced from nitrates. 27. Xanthomonas papavericola. . ccc. Ammonia formed in nitrate media. 28. Xanthomonas alfalfae. bbb. Starch not hydrolyzed. c. Nitrites produced from nitrates. 29. Xanthomonas acernae. cc. Nitrites not produced from nitrates. 30. Xanthomonas carotae. 31. Xanthomonas hederae. 32. Xanthomonas phormicola. 25. Xanthomonas vesicatoria. ccc. Ammonia formed in nitrate media. 33. Xanthomonas geranii. 152 MANUAL OF DETERMINATIVE BACTERIOLOGY bbbb. Starch hydrolysis not reported. c. Nitrites produced from nitrates. 34. Xanthomonas antirrhini. 35. Xanthomonas heterocea. cc. Nitrites not produced from nitrates. 36. Xanthomonas gummisudans . 37. Xanthomonas lactxicae. 38. Xanthomonas nigromaculans . aa. Gelatin not liquefied. b. Starch not hydrolyzed. 39. Xanthomonas oryzae. aaa. Gelatin not reported. b. Starch hydrolyzed. 40. Xanthomonas celebensis. 2. Colonies whitish to cream. a. Gelatin liquefied. b. Starch hydrolyzed. c. Nitrites produced from nitrates. 41. Xanthomonas panici. 42. Xanthomonas proteomaculans . 43. Xanthomonas majuhotis. cc. Nitrites not reported. 44. Xanthomonas ruhrisnhalhicans . bb. Starch not reported. 45. Xanthomonas cannae. 46. Xanthomonas zingiberi. 47. Xanthomonas conjaci. 1. Xanthomonas hyacinthi (Wakker) Milk: Casein is precipitated and Dowson. {Bacterium hyacinthi Wakker, digested. Tyrosine crystals produced. Botan. Centralblatt, 14-, 1883, 315; Nitrites not produced from nitrates. Bacillus hyacinthi Trevisan, I generi e le Indole : Slight production, specie delle Batteriacee, 1889 ; 19 ; Pseudo- Hydrogen sulfide is produced. monas hyacinthi Erw. Smith, Bot. Acid, no gas, from glucose, fructose, Gazette, ^4) 1897, 188; Phyiomonas galactose, suci'ose and maltose. hyacinthi Bergey et al., Manual, 1st ed., Starch: Hydrolysis slight. 1923, 177; Dowson, Cent. f. Bakt., II Optimum temperature 28° to 30°C. Abt., 100, 1939, 188.) From Gr. hy- Maximum 34° to 35 °C. Minimum4°C. acinthus, the hyacinth; M. L. Hi/a- Aerobic, with the exception of maltose, cinthus, a generic name. where it is facultative anaerobic. Description from Smitli, Div. Veg. Habitat: Produces a yellow rot of Phys. and Path., U. S. D. A. Bui. 26, hyacinth bulbs, Hyacinthus orientalis. 1901, 40. Rods: 0.4 to 0.6 by 0.8 to 2 microns. 2. Xanthomonas pruni (Erw. Smith) Motile with a polar flagellum. Filaments Dowson. {Pseudomonas pruni Erw. present. Gram-negative. Smith, Science, N. S. 17, 1903, 456; Gelatin: Slow liquefaction. jBacieHfwi pr?ider). Lipolytic (Starr and Burkholder, Phy- topath.,3;?, 1942, 600). Acid but not gas from glucose, fruc- tose, sucrose, lactose, galactose, glycerol and dextrin. Certain strains hydrolyze starch, others do not (Burkholder and Li, Phytopath., 31, 1941, 753). Optimum temperature 30°C. Source : Isolated from spotted tomato fruits in South Africa. Habitat : Pathogen on tomatoes, Lycu- persicon esculentum and peppers, Capsi- cum annuum. 25a. Xanthomonas vesicatoria var. raphani (White) Starr and Burkholder. {Bacterium vesicatoria var. raphani White, Phytopath., 20, 1930, 653; Phylo- monas vesicatoria var. raphani Burk- holder, in Manual, 5th ed., 1939, 154; Starr and Burkholder, Phytopath., 32, 1942, 600.) From M. L. Raphanus, the radish, a generic name. Distinctive characters : Cultural char- acters similar to Xanthomonas vesica- toria, but differs in that it is able to attack radishes, turnips, and other crucifers. Differs from Xanthomonas campestris in that it does not cause a vascular disease, and differs from Xanthomonas cainpestris var. armoraciae in that it is not patho- genic on horseradish. Source : Isolated from leaf spots of radish and turnips in Indiana. Habitat : Pathogenic on radish, turnips, and other crucifers ; and on tomato and pepper. 26. Xanthomonas nakatae (Okabe) Dowson. {Bacterium nakatae Type B, Okabe, Jour. Soc. Trop. Agr., Formosa, 5, 1933, 161 ; Phylomonas nakatae Burk- holder, in Manual, 5th ed., 1939, 154; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 12.) Named for Nakata, the Japa- nese plant pathologist. Rods: 0.3 to 0.4 by 1.1 to 2.5 microns. Capsules. Motile with a polar flagellum. Gram-negative. Gelatin: Liquefaction. Brown color. Beef -extract agar colonies: Amber yel- low, round, smooth, glistening, margins entire. Brown. Broth : Moderate turbidity with yellow ring. Medium turns brown. Milk: Casein is precipitated and di- gested. Tyrosine crystals. Brown color. Nitrites not produced from nitrates. Indole not formed. Slight amount H2S produced. Acid but not gas from glucose, sucrose, maltose and lactose. Starch: Strong diastatic action. Optimum temperature 30° to 32°C. Maximum 39°C. Minimum 10°C. No growth in beef extract broth plus 2 per cent salt. Aerobic. Distinctive character: Differs from Type A in that it produces a brown pig- ment in culture. (Description of Type A not seen.) Source : Isolated from water-soaked to brown leaf spots on jute. Habitat : Pathogenic on jute, Corchoriis capszdaris. 27. Xanthomonas papavericola (Bryan and McWhorter) Dowson. {Bacterium papavericola Bryan and McWhorter, Jour. Agr. Res., 40, 1930, 9; Phytomonas papavericola Bergey et al., Manual, 4th ed., 1934, 266; Dowson, Cent. f. Bakt., IIAbt.,iOO, 1939,190.) FromL. papaver, poppy; -cola, dweller; M. L. Papaver, a generic name. Rods: 0.6 to 0.7 by 1 to 1.7 microns. Chains. Capsules. Motile with a single polar flagellum. Gram-negative. FAMILY PSEUDOMONADACEAE 165 Gelatin : Liquefaction. Beef agar colonies : Mustard yellow to primuline yellow, circular, margins entire. Broth : Turbidity prompt with a yellow ring and an incomplete pellicle. Milk : Soft coagulation, peptonization and production of tyrosine crystals. Nitrates : A weak reaction for nitrites after 10 days. Indole not formed. Hydrogen sulfide is produced. Lipolytic (Starr and Burkholder, Phy- topath., 5:?, 1942, 600). Acid but not gas from glucose, galac- tose, fructose, sucrose, lactose, maltose, glycerol and mannitol. Starch is hydrolyzed. Optimum temperature 25° to W°C. Maximum 35°C. No growth in broth plus 5 per cent salt. Aerobic. Source: Isolated from black spots on leaves, buds and pods of poppy. Habitat : Pathogenic on poppy, Papaver rhoeas. 28. Xanthomonas alfalfae (Riker et al.) Dowson. {Bacterium alfalfae Riker, .Tones and Davis, Jour. Agr. Res., 51, 1935, 177; Phylomonas alfalfae Riker ot al., ihid.; Pseudomonas alfalfae Riker ot al., ihid.; Dowson, Trans. Brit. Alycol. Soc, 26, 1943, 11.) From Spanish, of alfalfa. Rods: 0.45 by 2.4 microns. Motile with a polar fiagellum. Gram-negative. Gelatin: Liquefied. Nutrient agar stroke : Growth abun- dant, filiform, smooth, glistening, butyr- ous, pale yellow. Broth: Turbid in 24 hours. Light sediment. Milk: Casein is precipitated and di- gested. Ammonia formed slowly in a nitrate medium. Carbohydrates : No acid in yeast broth plus sugars. Starch is hydrolyzed. Aerobic. Optimum temperature 24° to 32°C. Maximum below 36°C. Minimum below 4°C. Source : Six single cell cultures isolated from diseased alfalfa. Habitat : Pathogenic on the leaves of alfalfa, Medicago sativa. 29. Xanthomonas acernea (Ogawa) comb. nov. {Pseudomonas acernea Ogawa, Ann. Phyt. Soc. Japan, 7, 1937, 123; Phylomonas acernea Ark, Phytopath., 29, 1939, 968.) From L. acerneus, of the maple. Rods: 0.2 to 0.6 by 0.5 to 1.2 microns. Motile with one polar fiagellum. Gram- negative. Gelatin: Liquified. Agar colonies : Round, smooth, convex, white to citron yellow, glistening, trans- lucent with amorphous structure. Broth : Turbid. Milk: Slowly cleared, slightly acid. No coagulation. Nitrites produced from nitrates. Hydrogen sulfide produced. No gas produced in peptone water plus sugars. Starch not hydrolyzed. Optimum temperature about 32°C. Thermal death point 59°C. Aerobic. Source : From diseased leaves of Acer trifidum in Japan. Habitat : Causes a disease in Acer spp. and in Aesculus turbinaia and Koelren- teria panicvlata. 30. Xanthomonas carotae (Kendrick) Dowson. {Phylomonas carotae Ken- drick, Jour. Agr. Res., 49, 1934, 504; Pseudomonas carotae Kendrick, ibid. ; Dowson, Cent. f. Bakt., II Abt., 100, 1939, 190.) From L. carota, the carrot. Rods: 0.42 to 0.85 by 1.38 to 2.75 microns. Motile with 1 or 2 polar flagella. Gram-negative. Gelatin: Liquefied. Potato glucose agar: Colonies round, smooth, glistening, margins entire, straw yellow in color. 166 MANUAL OF DETERMINATIVE BACTERIOLOGY Milk : Casein precipitated and milk cleared; alkaline. Nitrites not produced from nitrates. Indole not formed. Acid, no gas, from glucose, d-galac- tose, xylose, rf-mannose, Z-arabinose, sucrose, lactose, raffinose, trehalose, d-mannitol and glycerol. No acid from maltose and rhamnose. Starch not hydrolyzed. Optimum temperature 25° to 30°C. Tolerates 4 per cent salt at pH 7. Aerobic. Source : Two original isolations from diseased carrots and a reisolation from inoculated carrots were used for the description. Habitat: Pathogenic on leaves of Daucus caroin var. saliva. 31. Xanthomonas hederae (Arnaud) Dowson. {Bacterium hederae Arnaud, Compt. rend. Acad. Sci., Paris, 171, 1920, 121; Phytomonas hederae Burk- holder and Guterman, Phj^topath., 22, 1932, 783; Dowson, Cent. f. Bakt., II Abt., 100, 1939, 190.) From L. hedera, ivy; M. L. Hedera, a generic name. Description taken from Burkholder and Guterman {loc. cit.). Rods: 0.6 by 2.13 microns. Motile with a single polar flagellum. Gram- negative. Gelatin: Liquefied. Beef-extract-agar slants: Growth good, filiform, amber yellow, butyrous. Broth: Turbid. Milk: Casein is precipitated and di- gested. Milk becomes alkaline. Nitrites not produced from nitrates. Hydrogen sulfide is formed. Indole not formed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 600). Acid from glucose, fructose, galactose, xylose, sucrose, lactose and glycerol. Alkali from salts of acetic, citric, lactic, malic and succinic; acids. The following are not utilized : arabinose, rhamnose, maltose, salicin, starch, cellulose and formic acid. Aerobic, facultative. Source : Isolated from diseased ivy leaves. Habitat : Pathogenic on ivy, Hedera helix. 32. Xanthomonas phormicola (Taki- moto) Dowson. {Bacterium phormicola Takimoto, Jour. Plant Protect., 20, 1933, 777; Phytomonas -phormicola Burkholder, in Manual, 5th ed., 1939, 159; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 12.) From M. L. Phormium, a generic name. Description translated by Dr. K. Togashi . Rods: 0.5 to 0.6 by 1 to 2 microns. Motile, with a single flagellum. Gram- negative. Gelatin : Liquefied. Agar colonies : Light yellow, then waxy yellow; butyrous, then viscid. Broth : Turbid, pellicle formed. Milk: Casein coagulated slowly and precipitated, then digested. Alkaline. Nitrites not produced from nitrates. Indole not formed. Hydrogen sulfide produced. No gas from sucrose, glucose, lactose and glycerol. No acid from various sugars in broth. Optimum temperature about 29°C. Maximum 39°C. Minimum about 0°C. Aerobic. Source : Species isolated from New Zealand flax, Phormium tenax. Habitat: Causes a leaf stripe of Phor- mium tenax. 33. Xanthomonas geranii (Burkholder) Dowson. {Phytomonas geranii Burk- holder, Phytopath., 27, 1937, 560; Dow- son, Cent. f. Bakt., II Abt., 100, 1939, 190.) From Greek, geranos, crane; M. L. Geranium , a generic name. Rods: 0.75 to 2.0 microns. Motile with a single polar flagellum. Gram- negative. Gelatin: Liquefied. Beef -extract agar slants : Moderate to good filiform growth, glistening, primu- line yellow. Develops in 24 hours. FAMILY PSEtJDO:MONADACEAE 167 Broth : Turbid in 24 hours. No pellicle but a moderate sediment. Milk : Becomes clear with a heavy casein precipitate. Peptonization with crystal formation. Nitrates reduced to ammonia. Indole not formed. Hydrogen sulfide formed. Lipolytic (Starr and Burkholder, Phy- topath., 32, 1942, 600). Acid from glucose, galactose, fructose, xylose, rhamnose, lactose, sucrose, raf- finose and glycerol. Alkaline reaction from salts of citric, malic, malonic and succinic acid. No growth in arabinose or formic, hippuric, maleic or tartaric acid. Starch not hydrolyzed. Aerobe. Distinctive characters : Pathogenic on Geranium spp., not on the house gera- nium, Pelargonium hortorum. In culture similar to Xanlhomonas pelargonii. Source : Three cultures isolated from Geranium- sangnineurn. Habitat : Pathogenic on Geranium san- guineum, G. maculatum, G. pratense and G. sylvaticum. 34. Xanthomonas antirrhini (Taki- moto) Dowson. {Psendomonas anti- rrhini Takimoto, Bot. Mag. Tokyo, 34, 1920, 257; Bacterium antirrhini Elliott, Man. Bact. Plant Path., 1930, 93; Phyto- monas antirrhini Magrou, in Hauduroy et al., Diet. d. bact. path., Paris, 1937, 331; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 11.) From Gr. antirrhinum, snapdragon; M. L. Antirrhinum, a generic name. Description from Elliott (loc.cit.). Rods: 0.3 to 0.4 by 0.8 to 1.2 microns. Motile with polar flagella. Capsules. Gram-negative. Gelatin: Liquefied. Agar colonies: Round, glistening, white, later yellow. Milk: Coagulated and casein digested. Nitrites are produced from nitrates. No gas produced. Aerobic. Optimum temperature 26° to 27°C. Maximum 34°C. Habitat: Causes a leaf spot of Anti- rrhinum majus. 35. Xanthomonas heterocea (Vzoroff) comb. nov. {Phytomonas heterocea Vzoroff, Bull. North Caucasian Plant Prot. Sta. Roztoff-on-Don, 6-7, 1930, 263; Bacterium heteroceum Burgwitz, Phytopathogenic bacteria, Leningrad, 1935, 135.) From Gr. hcterus, another, different. Description taken from Rev. App. Myc, 10, 1931, 628. Rods : 0.4 to 0.6 by 1.0 to 2.0 microns. Motile. Gram-negative. Gelatin : Slow liquefaction. Agar colonies : Round, convex, smooth, semi-transparent, glistening, yellow to amber, 2 mm. in diameter. Pitted sur- face . Milk: No coagulation. At first acid, later alkaline. Nitrites produced from nitrates. Indole not formed. Hydrogen sulfide produced. Acid from glucose, galactose, arabi- nose, xylose, sucrose, maltose, salicin, glycerol and mannitol . Does not ferment lactose, inulin, ethyl alcohol, esculin, adonitol or dulcitol. Optimum temperature 25° to 30°C. Source : Isolated from diseased tobacco in the North Caucasus. Habitat : Pathogenic on Nicotiana tabacum. 36. Xanthomonas gummisudans (Mc- Culloch) Starr and Burkholder. (Bac- terium gummisiulans McCulloch, Phyto- path., H, 1924, 63; also Jour. Agr. Res., 27, 1924, 229; Pseudomonas gummisudans Stapp, in Sorauer, Handb. d. Pflanzen- krank., 2, 5 Aufl., 1928, 54; Phytomonas gummisudans Bergey et al., Manual, 2nd ed., 1925, 201; Starr and Burkholder, Phytopath., 32, 1942, 600.) From L. gummi, gum; sudans, sweating, dripping. Rods : 0.6 to 0.8 by 1 to 2.8 microns. 168 MANUAL OF DETERMINATIVE BACTERIOLOGY Capsules. Motile with a polar flagellum. Gram-negative. Gelatin : Liquefied. Beef -peptone agar colonies : Amber yellow, circular, transparent, smooth, with definite margins. Broth : Moderately turbid with a yel- low ring. Milk : Soft curd which is digested with formation of tyrosine crystals. Nitrites not produced from nitrates. Indole not produced. Hydrogen sulfide produced. Lipolytic (Starr and Burkholder, loc. cit.). Acid from glucose and sucrose. Optimum temperature 30°C. Maxi- mum Se'C. Minimum 2°C. Aerobic. Source : From gummy lesions on gladio- lus leaves. Habitat : Pathogenic on leaves of glad- ioli. 37. Xanthomonas lactucae (Yama- moto) Dowson. (Bacterium lactucae Yamamoto, Jour. Plant Protect., 21, 1934, 532; Phytomonas lactucae Bergey et al.. Manual, 5th ed., 1939, 163; Dow- son, Trans. Brit. Mycol. Soc, 26, 1943, 12.) From L. lactuca, lettuce; M. L. Lactuca, a generic name. Description translated by Dr. K. Togashi . Rods : 0.6 to 0.8 by 1.75 to 2.8 microns. Motile with a polar flagellum. Gram- negative. Gelatin : Liquefaction slow. Agar colonies: Circular, convex, mar- gin entire, surface smooth, wet-shining, yellow. Broth: Turbid. Ring and pellicle. Milk: Slow peptonization. Nitrites not produced from nitrates. Indole not produced. Hydrogen sulfide produced. Acid, no gas, from glucose, sucrose, and lactose in bouillon; no acid from glycerol in bouillon. Optimum temperature 28°C. Maxi- mum Z5°C. Minimum below 2''C. Aerobic. Source : Isolated from leaf spot of lettuce. Habitat : Pathogenic on leaves of as- paragus lettuce, Lactuca saliva var. an- gustata. 38. Xanthomonas nigromaculans (Takimoto) Dowson. (Bacterium nigra - m.aculans Takimoto, Jour. Plant Protect., Tokyo, H, 1927, 522; Phytomonas nigro- maculans Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 387; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 12.) From L. niger, black; macu- lans, spotting. Description translated by Dr. K. Togashi . Rods: 0.6 to 0.9 by 1.5 to 2.8 microns. Motile with 1 or 2 polar flagella. Gram- negative. Gelatin: Liquefaction. Agar colonies : Yellow, circular, margins entire, smooth, glistening. Broth : Growth moderate with yellow pellicle. Milk : Coagulation and digestion of the casein. Nitrites not produced from nitrates. Indole not produced. No acid or gas from glucose, sucrose, lactose, mannitol and glycerol in peptone water. Optimum temperature 27° to 28°C. Maximum 33°C. Minimum O^C. Aerobic. Source : Isolated from lesions on leaf and petioles of burdock. Habitat: Pathogenic on leaves and petioles of Arctium lappa, the burdock. 39. Xanthomonas oryzae (Uyeda and Ishiyama) Dowson. {P seudomonas oryzae Uyeda and Ishiyama, Proc. Third Pan-Pacific Sci. Congr., Tokyo, 2, 1926, 2112; Bacterium- oryzae Nakata, see Elliott, Man. Bact. Plant Path., 1930, 172; Phytomonas oryzae Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 388; Dowson, Trans. Brit. FAMILY PSEUDOMONADACEAE 169 Mycol. Soc, 26, 1943, 12.) From Gr. oryza, rice; M. L. Oryza, a generic name. Probable synonym : Pseudomonas ito- ana Tochinai, Ann. Phytopath. Soc. .Japan, 2, 19.32, 456; Bacterium itoanum Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 74; Phytomonas itoana Magrou, in Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 370. Rods: 0.5 to 0.8 by 1.0 to 2.0 microns. Motile with a polar flagellum. Gram- negative. Gelatin: No liquefaction. Nutrient agar colonies : Round, smooth, glistening, wax yellow. Milk: Slightly acid. Nitrites are not produced from nitrates. Hydrogen sulfide produced. Acid but no gas from glucose, lactose and sucrose. Optimum temperature 26° to 30°C. Strict aerobe. Source : Isolated from a leaf blight of rice. Habitat : Pathogenic on rice, Oryza saliva. 40. Xanthomonas celebensis (Gau- mann) Dowson. {Pseudomonas celeben- sis Gaumann, Ztschr. f. Pflanzenkrank., S3, 1923, 11; Meded. Inst, voor Planten- ziek., Buitenzorg, 59, 1923, 17; Bacterium celebense Elliott, Man. Bact. Plant Path., 1930, 108; Phytomonas celebensis Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 343; Dowson, Trans. Brit. Mycol. Soc, 26, 1943, 11.) From M. L. of the island Celebes. Rods : 0.9 by 1.5 microns. Motile by a polar flagellum. Gram -negative. Agar colonies: Grayish yellow. Broth: Thin pellicle. Milk: Coagulated and cleared. Nitrites not produced from nitrates. Sodium selenite: Brick red. Starch is hydrolyzed. Source : From vascular bundles of diseased bananas in Celebes. Habitat : Causes the blood disease of banana. 41. Xanthomonas panici (Elliott) comb. nov. {Bacterium panici Elliott, Jour. Agr. Res., 26, 1923, 157; Pseudo- monas panici Stapp, in Sorauer, Handb. d. Pflanzenkrank., 2, 5 Aufl., 1928, 27; Phytomonas panici Bergey et al., Man- ual, 3rd ed., 1930, 269.) From M. L. Panicum, a generic name. Rods : 0.69 by 1.66 microns. Capsules. Motile by 1 or rarely 2 polar flagella. Gram-negative. Gelatin : Liquefaction slow. Beef agar colonies: Round, white, smooth, glistening, margins at first entire, later undulate. Broth : Moderate turbidity in 24 hours. Thin pellicle. Medium brownish. Milk: Alkaline and clears. Nitrites are produced from nitrates. Indole not produced. Hydrogen sulfide produced. No gas from carbohydrates. Starch: Hydrolysis moderate. Optimum temperature 33°C. Maxi- mum 45°C. Minimum 5°C. Optimum pH 6.15 to 6.3. pH range 5.4 to 10.0. Aerobic. Distinctive characters : Differs from Pseudomonas andropogoni in that it liquefies gelatin, produces nitrites from nitrates, and does not infect sorghum and broom corn. Source : Isolation from water soaked lesions on leaves, sheaths and culms of millet collected in Wisconsin and in S. Dakota. Habitat : Pathogenic on proso millet, Panicum m.iliaceum, 42. Xanthomonas proteamaculans (Paine and Stansfield) comb. nov. {Pseu- domonas proteamacidans Paine and Stans- field, Ann. Appl. Biol., 6, 1919, 38; Phytomonas proteamacidans Bergey et al.. Manual, 3rd ed., 1930, 247; Bacterium proteamacidans Elliott, Man. Bact. Plant Path., 1930, 186.) From M. L. Protea, a generic name; maculans, spotting. Rods : 0.6 to 0.8 by 0.8 to 1.6 microns. 170 MANUAL OF DETERMINATIVE BACTERIOLOGY Motile with 1 to 3 polar flagella. Gram- positive. Gelatin : Liquefaction. Agar slant : Growth wet-shining, dirty white with a faint yellow tinge. Broth: Turbid in 24 hours. Slight ring. Milk : Acid with soft curd after 2 days. Later a separation of whey. Nitrites are produced from nitrates. Acid and gas from glucose, sucrose and mannitol. No acid or gas from lactose. Starch: Slight hydrolysis. Source : Repeated isolation from a leaf- spot of Proica in England. Habitat : Pathogenic on Protea cy- nar aides. 43. Xanthomonas manihotis (Arthaud- Berthet) comb. nov. {Bacillus manihotus Arthaud-Berthet by Bondar, Chacaras and Quintaes 5(4), 1912, 15; Bacillus manihot Bondar (and Arthaud-Berthet), Bol. Agric, Sao Paulo, 16, 1915, 513; Bacterium m.anihoius Drummond and Hipolito, Ceres, 2, 1941, 298; Phytomonas manihotis Viegas, Rev. d. Agr., Pieraci- caba, 15, 1940, 475.) From M. L. Mani- hotus, a generic name. Description from Burkholder, Phyto- path., 32, 1942, 147. Rods : 0.35 to 0.93 by 1.4 to 2.8 microns. Gram-negative and mostly non-motile. One isolate showed a few cells with 1 polar flagellum. Amaral (Instit. Biol., Sao Paulo, Arq., IS, 1942, 120) states that the species is motile with one polar fla- gellum. Gelatin: Liquefaction. Beef -extract-peptone agar : Streaks raised, ivory-color, smooth, shiny, with edges entire. Potato-glucose agar : Growth abun- dant, white to hyaline, very mucoid. Broth : Turbid with a whitish granular ring. Litmus milk : Litmus reduced and milk clears. With return of color, litmus is purple. Indole not formed. Hydrogen sulfide is formed. Nitrites produced from nitrates (Drummond and Hipolito, loc. cit.). Asparagine not used as a nitrogen and carbon source. No growth in nitrate synthetic broth. Weak growth but slight acid production in synthetic medium plus glucose, d-galactose, d-fructose, d-xylose, maltose and sucrose. No growth in rhamnose, 1-arabinose, d-lactose, glycerol, mannitol and salicin. Good growth with alkaline reaction in same medium plus salts of the following acids: acetic, citric, malic, maleic and succinic. The salts of formic, hippuric, lactic and tartaric acids were not utilized. Starch not hydrolyzed. Amaral {loc. cit.) finds hydrolysis. Lipolytic action slight. Aerobic. Optimum temperature 30°C. Maxi- mum 38°C. Minimum 5°C. Source : First isolated from the cassava, Manihotus utilissima in Brazil. Habitat : Produces a wilt disease on various species of Manihotus. 44. Xanthomonas rubrisubalbicans (Christopher and Edgerton) comb. nov. {Phytomonas rubrisubalbicans Christo- pher and Edgerton, Jour. Agr. Res., 4^, 1930, 266; Bacterium rubris2tbalbicans Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 105.) From L. ruber, red; subalbicans, nearly white. Rods: Short with polar flagella. Cap- sules. Gram-negative. Gelatin: No liquefaction. Bacto-glucose agar colonies: Circular, glistening, viscid, milky gray to buff. Margins translucent, entire. Broth : Turbid after 24 hours. Pellicle and a ropy sediment. Indole produced. Hydrogen sulfide produced. No gas from carbohydrates. Starch hydrolyzed. Optimum temperature 30°C. Optimum pH 6.8 to 8.0. Source : Isolated many times from mot- tled stripe of sugar cane in Louisiana. FAMILY PSEUDOMOXADACEAE 171 Habitat : Pathogenic on sugar cane, Johnson's grass and sorghum. 45. Xanthomonas cannae (Bryan) comb. nov. {Bacterium cannae Bryan, Jour. Agr. Res.,^i, 1921, \b2;Phytomonas cannae Bergey et al., Manual, 1st ed., 1923, 188; Pseudomonas cannae Stapp, in Sorauer, Handb. d. Pflanzenkrank., 2, 5 Aufl., 1928, 65.) From Gr. canna, a reed; M. L. Canna, a generic name. Rods: 0.5 to 0.7 by 1.0 to 2.0 microns. Motile with 1 to 3 polar flagella. Cap- sules. Gram -negative. Gelatin: Slow liquefaction. Agar streaks: Filiform, white, moist, with thin margins and granular centers. Broth: Turbid, heavy sediment. Milk: Alkaline and clears. Nitrites are produced from nitrates. Indole not produced. Hydrogen sulfide produced. Optimum temperature 35°C. Maxi- mum 40°C. Minimum 5°C. Aerobic. Source : Isolated from diseased canna leaves collected in Washington, D. C. and in Illinois. Habitat : Causes disease in Canna indica. 46. Xanthomonas zingiberi (LVeda) comb. nuv. (Uyeda, Cent. f. Bakt., II Abt., 17, 1907, 383; Pseudomonas zingi- beri Uyeda, Rept. Imp. Agr. Exp. Sta., Japan, No. 35, 1908, 114; Bacterium zingiberi Nakata, see Elliott, Man. Bact. Plant Path., 1930, 266; Phytomonas zingi- beri Magrou, in Hauduroy, et al., Diet. d. Bact. Path., Paris, 1937, 437.) From L. zingiberis, ginger; M. L. Zingiber, a generic name. Description from Stapp, in Sorauer, Handb. d. Pflanzenkrank., 2, 5 Aufl., 1928, 65. Rods : 0.5 to 1.1 by 0.75 to 1.8 microns. Non-motile at first, later a polar flagel- lum. Gram-negative. Gelatin: Liquefaction. Agar colonies : White. Milk: Coagulation and peptonization of the casein. Nitrites are produced from nitrates. Indole not formed. Hydrogen sulfide is formed. No gas from glucose. Optimum temperature 28°C. Maxi- mum 40°C. Minimum 5°C. Source : Isolated from ginger plant showing a rot at the base of the sprouts. Habitat: Pathogenic on ginger, Zingi- ber officinale. 47. Xanthomonas conjaci (Uyeda) comb. nov. (Pseudomonas conjac Uyeda, Bot. Mag. Tokyo, 24, 1910, 182; Bac- terium conjac Elliott, Man. Bact. Plant Path., 1930, 121; Phytomonas conjac Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 347.) From M. L. conjac, the specific name of the plant which this species attacks. Description from Elliott (loc.cit.). Rods: 0.75 to 1.0 by 1.5 microns. Motile with 1 to 4 polar flagella. Gram- positive. Gelatin colonies : Circular to irregular, light yellow. Broth: Pellicle formed. Milk: Coagulation. Conjac: Liquefied. Nitrites produced from nitrates. Indole produced. Hydrogen sulfide produced. Gas from glucose. Favorable temperature 24°C. Habitat : Pathogenic on Amorphophal- lus konjac. Appendix I:* The following organisms placed in the genus Psei^domottas apparently belong in Xanthomonas. Some may even be plant pathogens although they were * Prepared by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, July, 1943. 172 MANUAL OF DETERMINATIVE BACTERIOLOGY isolated from water, soil and similar sources. Pigment is usually yellow and is not water-soluble. Key to yellow and other chromogenic species in genus Pseudomonas. 1. Colonies yellow. a. Gelatin liquefied. b. Nitrites produced from nitrates. c. Acid and gas produced from glucose. 1 . Pseudomonas fermentans . cc. Acid but no gas from glucose. 2. Pseudomonas trifolii. 3. Pseudomonas xanthe. ccc. Action on glucose not recorded. 4. Pseudomonas caudata. bb. Nitrites not produced from nitrates. c. Litmus milk acid or ferment lactose. 5. Pseudomonas perlurida. 6. Pseudomonas iridescens. cc. Litmus milk not coagulated. Yellow sediment. 7. Pseudomonas turcosa. ccc. Litmus milk slimy, alkaline. 8. Psezidomonas ochracea. aa. No liquefaction of gelatin. b. Nitrites produced from nitrates. c. Litmus milk, slow coagulation. 9. Pseudomonas cerevisiae. cc. Litmus milk, acid but no digestion. 10. Pseudomonas arguta. ccc. No growth in litmus milk. 11. Pseudomonas subcreta. cccc. Action on litmus milk, not recorded. 12. Pseudomonas pictorjun. bb. Nitrites not produced from nitrates. c. Butter colored pellicle on litmus milk. 13. Pseudomonas lacunogenes. cc. No surface pellicle. 14. Pseudomonas segnis. 2. Colonies on gelatin blue center surrounded by yellow zone with peripheral green zone, a. Gelatin liquefied. b. Nitrites produced from nitrates. 15. Pseudomonas lemonnieri. 1. Pseudomonas fermentans von Wol- with rounded ends, occurring singly and zogen Kiihr. (von Wolzogen Kiihr, in pairs. Motile, with a single or occa- Cent. f. Bakt., II Abt., 85, 1932, 228; sionally 2 or 3 polar flagella. Gram- Flavobacterium fermentans Bergey et al., negative. Manual, 4th ed., 1934, 155.) From Gelatin colonies: Circular, grayish, Latin, fermento, to ferment. with rapid liquefaction. Rods: 0.4 to 0.6 by 1.7 to 3.4 microns. Gelatin stab : Liquefaction crateriform. FAMILY PSEUDOMONADACEAE 173 Agar colonies : Circular, slightly con- vex, opaque, gray by reflected, and light-brown by transmitted light. Agar slant : Gray, becoming yellowish. Broth: Turbid with pellicle. Litmus milk: Acid. Potato : Gray to yellowish growth. Indole is formed. Nitrites produced from nitrates. Acid and visible gas from glucose, lactose and sucrose. Acetylmethylcarbinol is formed. Ammonia is formed from peptone and asparagin. Hydrogen sulfide is formed. Starch is hydrolyzed. Lipase is formed. Catalase positive. Aerobic, facultative. Optimum temperature 37°C. Distinctive character; Produces gas in lactose fermentation tubes. Source : Ten cultures from the larvae of a midge (Chironomtis plumosus) and from filtered water. Habitjit: Unknown. 2. Pseudomonas trifolii Huss. (Huss, Cent. f. Bakt., II Abt., 19, 1907, 68; Flavobacterium trifolii Bergey et al., Manual, 1st od., 1923, 111.) From Latin, tres (Iri-), three; Jolivm, leaf; M. L. Trifolium, clover. Possible synonym : Bacillus annulatus Wright. (Wright, Memoirs Nat. Acad. Sci., 7, 1895, 443; Pseudomonas annidata Chester, Man. Determ. Bact., 1901, 315; Relationship to Bacillus annulatus Zim- mermann uncertain. Die Bakt. unserer Trink- und Nutzwiisser, Chemnitz, II Reihc, 1890, 30; Flavobacterium annula- lum Bergey et al., Manual, 1st ed., 1923, 110.) According to Mack (Cent. f. Bakt., II Abt., 95, 1936, 218) the following organism is to be regarded as identical with Pseudo- monas trifolii: Bacillus mesentericus aureus Winkler (Cent. f. Bakt., II Abt., 5, 1899, 577) regarded by Burri (Cent. f. Bakt., II Abt., 10, 1902, 756) and Dtiggeli (Cent. f. Bakt., II Abt., 12, 1904, 602) as identical with the organism which Duggeli (loc. cit.) names Bacterium herbicola aureum. The organism studied as Bacterium herbicola by Hiittig (Cent, f. Bakt., II Abt., 84, 1931, 231) is not regarded as identical with the Burri and Diiggeli organism by Mack. Beijerinck (Cent. f. Bakt., II Abt., IB, 1905, 366) states that Bacillus herbicola of Burri and Diiggeli is identical with his Bacillus anglomerans (Botan. Ztg., 1888, 749). If so, this binomial has priority. Rods : 0.5 to 0.7 by 0.75 to 2.0 microns, occurring singly, in pairs and in chains. Motile, possessing a single polar flagellum. Gram-negative. Gelatin colonies : Convex, smooth, moist, glistening, grayish -yellow. Gelatin stab: Napiform liquefaction. Agar colonies: Small, circular, grayish, becoming brownish-yellow. Agar slant : Yellowish, becoming brown- ish-yellow streak, lacerate margin. Broth: Turbid, with grayish-yellow pellicle and sediment. Litmus milk: Slowly coagulated; alka- line; with yellow ring. Potato: Thick, yellowish, fiat, smooth, glistening. Hj'drogen sulfide produced. Indole is formed. -\cid from glucose, sucrose, xylose, arabinose, and mannitol. No acid from lactose. Nitrites produced from nitrates. Cultures have an agreeable odor. Volutin formed. Aerobic, facultative. Optimum temperature 33° to 35°C. Source: Isolated from clover hay. Habitat : Evidently a common organism on the leaves of plants. 3. Pseudomonas xanthe Zettnow. (Zettnow, Cent. f. Bakt., I Abt., Orig., 77, 1915, 220; Flavobacterium zettnowii Bergey et al.. Manual, 1st ed., 1923, 112; Flavobacterium xanthium (sic) Ber- gey et al.. Manual, 3rd ed., 1930. 145.) From Gr. xanthus, yellow. 174 MANUAL OF DETERMINATIVE BACTERIOLOGY Rods: 0.5 to 0.6 by 0.4 to 1.4 microns. Motile, possessing a single or occasionally two or more very long (20 microns) polar flagella. Gram-negative. Gelatin colonies: Circular, yellow, granular. Gelatin stab : Pale-yellow surface growth. Brownish yellow under surface colonies. Saccate liquefaction. Agar slant : Dark yellow, glistening, with dark yellow sediment in water of condensation. Pigment not water-sol- uble. Broth: Turbid. Litmus milk: Slightly acid. Litmus reduced. Potato : Grayish yellow to brownish growth. Indole formed. Nitrites are produced from nitrates. Acid formed in glucose. Starch hydrolyzed. Blood serum not liquefied. Aerobic, facultative. Optimum temperature 30°C. Source: Air contamination. 4. Pseudomonas caudata (Wright) Conn. {Bacillus caudatus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 444; Bacterium caudatus Chester, Annual Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 107; Conn, Jour. Agr. Res., 16, 1919, 313; Flavobacterium caudatum Bergey et al., Manual, 1st ed., 1923, 109.) From Latin, cauda, tail. Rods : Long, granular, slender, occurring singly, in pairs and in chains. Appear like cocci in old cultures. Motile, pos- sessing a polar flagellum (Conn). Gram- negative. Gelatin colonies: Yellow, translucent, smooth, undulate. Gelatin stab: Villous growth in stab. Crateriform liquefaction. Agar slant: Yellow to orange, glisten- ing, translucent, slightly spreading. May lose power to form pigment. Broth: Turbid, with yellow sediment. Litmus milk: Unchanged. Potato: Dark yellow, raised, rough, spreading. Indole not formed. Nitrites and ammonia produced from nitrates. Ammonia produced from peptone. Starch is digested. Aerobic, facultative. Optimum temperature 25°C. Habitat : Water. 5. Pseudomonas perlurida Kellerman et al. (Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 516; also McBeth, Soil Sci., 1, 1916, 472; Ccllulomonas perlurida Bergey et al., Manual, 1st ed., 1923, 163.) Rods: 0.4 by 1.0 micron. Motile with one to three polar flagella. Gram- negative. Gelatin stab: Liquefaction. Agar slant : Moderate, flat, faint yellow growth. Broth: Turbid in 5 days. Litmus milk : Acid. Peptonization after 16 days. Potato : Scant yellow growth with bleaching along line of growth. Indole not formed. Nitrites not produced from nitrates. Ammonia is produced. Acid from ghicose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source : Soil from Virginia, Louisiana and Missouri . Habitat: Soil. 5a. Pseudomonas perlurida var. vir- giniana Kellerman et al. {loc. cit.). Does not grow on potato and liquefies gelatin rapidly. Source : Soil from Virginia. 6. Pseudomonas iridescens Stanier. (Jour. Bact., 43, 1941, 542.) From Latin, iridescent. Rods: 0.2 to 0.3 by 1.5 to 7.0 microns, average length 5.0 to 6.0 microns, occur- ring singly. Non-motile. Gram-nega- tive. FAMILY PSEUDOMOXADACEAE 175 Sea water gelatin stab : Filiform growth. Liquefaction by some strains. Sea water agar colonies: Concave, 2 to 3 mm in diameter, smooth, glistening, translucent, pale yellow, edge irregular. After 2 to 3 days a marked iridescence. Later colonies rough, opaque, bright yellow, sunken central portion with trans- lucent periphery. Sea water agar slant : Growth spreading, smooth, glistening, translucent, pale yellow, iridescent, butyrous. Sea water broth: Turbid, light yellow, granular pellicle. Indole not formed. Nitrites not produced from nitrates. Hydrogen sulfide not produced. Catalase positive. Urease negative. Acid from xylose, glucose, galactose, /actose, maltose, sucrose and cellobiose. No acid from arabinose. Starch and cellulose are attacked. Aerobic. Optimum temperature 23°C. Mini- mum o°C. Maximum 30°C. Salt range : 0.25 to 6.0 per cent. Opti- mum 1.0 to 4.0 per cent. Source : Sea water. Habitat : Common along the coast of the North Pacific. 7. Pseudomonas turcosa (Zimmer- mann) Migula. {Bacillus turcosa Zim- mermann, Bakt. unserer Trink- und Nutzwasser, Chemnitz, 2, 1894, 32; Migula, Syst. d. Bakt. 2, 1900, 937; Flavo- bacteriiim turcosuvi Bergej^ et al., [Man- ual, 1st ed., 1923, 111.) From :\r. L. turcois, turquoise. Rods: 0.5 by 1.05 to 1.82 microns, oc- curring singly. A short polar fiagellum (Migula). Gram-negative. Gelatin colonies: Small, translucent, yellow. Gelatin stab: Small, yellow, convex surface growth, with slight brownish tint. Liquefaction, with grayish to greenish color in liquefied portion. Agar slant: Abundant, glistening, greenish to sulfur yellow streak. Broth: Slightly turbid with yellow sediment. Litmus milk: No coagulation. Yellow sediment. Potato : Clear chromium yellow growth over entire surface. Indole is not formed. Nitrites not produced from nitrates. Acid from glucose. Slight action on sucrose. Aerobic, facultative. Optimum temperature 30''C. Source : Isolated by Tataroff from a well in Dorpat (Die Dorpaten Wasser- bakterien, Inaug. Diss., 1891, 52, No. 24). Habitat : Water, sea water. 8. Pseudomonas ochracea (Zimmer- mann) Chester. {Bacillus ochraceus Zimmermann, Bakt. unserer Trink- und Nutzwasser, Chemnitz, 1, 1890, 60; Chester, Determinative Bacteriology, 1901, 316; Flavohacteritnn ochraceum Ber- gey et al., Manual, 1st ed., 1923, 110; Chromobacterium ochraceum Topley and Wilson, Princ. Bact. and Immun., 1, 1931, 405.) From Greek, ochros, pale yellow. Rods : 0.7 to 0.8 by 1.2 to 4.5 microns, occurring in pairs and longer chains. Slow undulatory motion (Zimmermann). Polar flagella (Lehmann and Neumann, Bakt.Diag.,lAufl.,^, 1896,255). Gram- negative. Gelatin colonies : Pale yellow to golden, ochre yellow, slightly raised, with slightly fringed margin, granular. Gelatin stab : Yellowish to yellow-gray surface growth. Infundibuliform lique- faction. Pale yellow to ochre yellow sediment. Agar colonies: Thin, flat, j^ellowish, smooth. Agar slant: Thin, yellowish-gray to ochraceous growth. Broth: Slightly turbid, with pale yel- low sediment. Litmus milk: Medium becomes slimy; alkaline. Potato : Ochre-yellow streak. Indole is formed. 176 MANUAL OF DETERMINATIVE BACTERIOLOGY Nitrites not produced from nitrates. Hydrogen sulfide is formed. Aerobic, facultative. Optimum temperature 35°C. Source: Chemnitz tap water. Habitat: Water. 9. Pseudomonas cerevisiae Fuhr- mann. (Fuhrmann, Cent. f. Bakt., II Abt., 16, 1906, 309; Flavohacterium cerevisiae Bergey et al., Manual, 1st ed., 1923, 111.) From Latin, cerevisia, beer. Rods: Straight and slightly curved, 0.6 by 1.5 to 2.0 microns, occurring singly and in chains. Motile, possessing tuft, four to six polar fiagella. Gram-negative . Gelatin colonies: Circular, white, slightly contoured, becoming brownish- yellow. Gelatin stab: Slight yellowish growth in stab. No liquefaction. Agar colonies: Thin, spreading, con- toured. Agar slant : Moist, glistening, thin, pale yellow, spreading, contoured. Litmus milk: Slow coagulation. Potato : Yellowish-brown, spreading growth. Indole not formed. Nitrites produced from nitrates. No gas from glucose . Aerobic, facultative. Optimum temperature 30°C. Source: Isolated from beer. Habitat: Unknown. 10. Pseudomonas arguta McBeth. (McBeth, Soil Science, 1, 1916, 465; Cellulomonas arguata (sic) Bergey et al., Manual, 1st ed., 1923, 164.) From Latin, arguo, to show. Rods: 0.3 by 0.8 micron. Motile with one or two polar fiagella. Gram-negative. Gelatin stab: Moderate, yellowish growth. No liquefaction in 30 days. Agar colonies: Circular, slightly con- vex, soft, grayish-white, granular, entire. Agar slant: Scant, grayish-white growth. Potato agar slant: Moderate, yellow- ish, glistening. Broth: Turbid. Ammonia cellulose agar: Enzymatic zone 2 to 3 mm in 30 days. Filter paper broth : Paper is reduced to loose fiocculent mass which disinte- grates very readily on slight agitation. More rapid decomposition when the broth contains ammonium sulfate, po- tassium nitrate, peptone or casein as sources of nitrogen. Litmus milk : Acid, not digested. Potato: No growth. Indole not formed. Nitrites produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, starch. No acid from glycerol, manni- tol or sucrose. Aerobic, facultative. Optimum temperature 20°C. Source: Isolated twice from California soils. Habitat: Soil. 1 1 . Pseudomonas subcreta McBeth and Scales. (McBeth and Scales, Bur. Plant Industry, U. S. Dept. Agr., Bui. 266, 1913, 37; Cellulomonas subcreta Bergey et al., Manual, 1st ed., 1923, 164.) From Latin, sub, under, imperfect; creta chalk. Rods : 0.3 by 1.4 microns. Motile with one to five polar fiagella. Gram-negative. Gelatin stab : Filiform growth, no lique- faction. Cellulose agar: No surface growth. Moderate, generally faint yellow growth in medium, area of growth sunken. Agar slant: Glistening, smooth, moisl., vitreous to faint yellow. Starch agar : Enzymatic zone 2 to 4 mm. Broth : No growth. Litmus milk : No growth. Potato : Growth scanty, concave due to slight liquefaction, white to faint yellow. Bleached around growth. Indole not formed. Tr^ce of nitrites produced from ni- trates. Ammonia not produced. Acid from glucose, lactose, maltose, FAMILY PSEUDOMONADACEAE 177 sucrose and starch. No acid from glyc- erol or mannitol. Aerobic, facultative. Optimum temperature 20°C. Habitat : Soil. 12. Pseudomonas pictorum Gray and Thornton. (Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 89; Achromobacter pictorum Bergey et al., Manual, 3rd ed., 1930, 217.) From Latin, picti, the Pict.s of Eastern Scot- land. Rods: 0.5 to 0.8 by 1.5 to 5.0 microns. Motile usually with a single polar flagel- lum. Gram-negative. Gelatin colonies : Circular, greenish- yellow, convex, smooth, glistening, en- tire. Gelatin stab: Xo liquefaction. Agar colonies : Circular, j^ellow, convex, smooth, glistening, entire. Agar slant: Filiform, yellow, convex, smooth, glistening, entire. Broth: Turbid. Nitrites produced from nitrates. Starch not hydrolj^zed. Acid from glucose and maltose. Attacks phenol. Aerobic, facultative. Optimum temperature 25''C. Source: One culture from soil. Habitat : Soil. 13. Pseudomonas lacunogenes Gor- esline. (Jour. Bact., 26, 1033, 447.) From Latin lacuno, dimple and gcnero, to produce. Short rods: 0.2 to 0.3 by 1.0 to 1.2 microns, with pointed ends, occurring singly or in pairs. Motile with a single polar flagellum from 2 to 15 microns in length. Gram-negative. Plain gelatin stab : No growth. Nutrient gelatin stab : Growth brown- ish-yellow, half-way down stab, heavier at surface. No liquefaction. Nutrient agar colonies: Small, yellow; surface of the agar pitted or dimpled. After 5 days colonies 5 to 7 mm in diam- eter, orange-yellow, slightly raised, sur- rounded by a depression. Nutrient agar slant: Growth heavy, light orange-yellow ; consistency of warm butter; edge entire, slightly raised. Shallow depression formed on each side of streak . Agar softened beneath growth . Nutrient broth: Turbid in 48 hours. Light orange-yellow pellicle ; considerable viscous sediment. Litmus milk: Alkaline; butter-colored pellicle. Reduction in bottom of tube after 10 days. No curd. No digestion. Potato: Growth moderate, orange-yel- low, smooth. No darkening. Indole not formed. Nitrites not produced from nitrates. Starch agar plates not hydrolyzed. Utilizes arabinose, galactose, lactose, fructose, maltose, melezitose, raffinose, starch, xylose, glucose, mannose, su- crose, pectin, rhamnose, salicin and dextrin. No growth in dulcitol, erythri- tol, glycerol, sorbitol, mannitol or inulin. Limits of pH : 5.4 to 10.0.' Temperature relations : Optimum 28''C. Good growth at 25°C. Moderate growth at 20° and at 37°C. No growth at 10° and at 42°C. Facultative anaerobe. Distinctive characters: Softens agar; considerable change in viscosity of agar due to this digestion; utilization of ammonium sulfate as nitrogen source. Source : Three cultures isolated from an experimental trickling filter receiving creamery wastes. Habitat : Probably widely distributed in nature. 14. Pseudomonas segnis Goresline. (Jour. Bact., 26, 1933, 452.) From Latin segne, non-energetic. Short rods: 0.2 to 0.3 by 1.0 to 1.2 mi- crons, with pointed ends, occurring singly or in pairs. Motile with a single polar flagellum. Gram-negative. Plain gelatin stab: No growth. Nutrient gelatin stab: Growth yellow, half-way down stab, best at surface. No liquefaction. 178 MANUAL OF DETERMINATIVE BACTERIOLOGY Nutrient agar colonies: Very small, light yellow; surface pitted. After 5 days colonies 5 mm in diameter. Nutrient agar slant : Grovv'th heavy, orange-yellow, consistency of warm but- ter; edge entire, slightly raised; slight depression formed on each side of growth. Agar softened beneath growth. Nutrient broth: Turbid in 48 hours. No pellicle or surface growth. Moderate amount of sediment. Old cultures with a yellow ring at surface and occasionally a loose membrane. Litmus milk : Slightly alkaline after 10 days. No reduction. No surface growth. Potato: Scant yellow-orange growth. No darkening. Indole not formed. Nitrites not produced from nitrates. No HoS produced. Starch not hydrolyzed. Utilizes arabinose, glucose, galactose, lactose, fructose, maltose, mannose, xylose, sucrose, melezitose and raffinose. Limits of pH : 5.8 to 9.0. Temperature relations : Optimum 28°C. Good growth at 25°C. Moderate growth at 20° and at 37°C. No growth at 10° and at 42°C. Facultative anaerobe. Distinctive characters: Softens agar; considerable change in viscosity of agar due to this digestion. Source : Isolated from an experimental trickling filter receiving creamery wastes. Habitat : Probably widely distributed in nature. 15. Pseudomonas lemonnieri (Lasseur) comb. nov. (Bacillus lemonnieri Lasseur, Compt. rend. Soc. Biol. Paris, 74, 1913, 47; Bui. de la Soc. des Sci. de Nancy, 1924; Flavobacicrium lasseuri Bergey et al.. Manual, 3rd ed., 1930, 144.) Named for Prof. G. le Monnier, a French scientist. Rods: 0.5 to 0.7 by 1.0 to 2.0 microns, occurring singly and in pairs. Motile with a single polar flagellum. Gram- negative. Gelatin colonies (glucose) : Circular with blue center, a granular, yellow zone and a peripheral blue zone. Rapid liquefaction with blue crystals. Gelatin stab : Liquefied. Agar colonies: Circular, yellowish, lobate margin. Agar slant : Yellowish streak, smooth, glistening. Broth: Turbid with thin pellicle. Litmus milk: After 48 hours the sur- face of the milk becomes yellow to cream color turning blue. A soft coagulum is formed. Potato : Raised growth, Prussian blue in color, with variations. Indole is not formed. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 22° to 25°C. Habitat : Water. Appendix II:* The following inade- quately described species may belong to the genus Xanthomonas. Bacterium citri deliciosae Passalacqua. (Rev. Pat. Veg., 24, 1934, 27.) Isolated from Citrvs sp. Bacterium malvacearum var. harbadense Evelyn. (Ann. Rept. Agric. Barbados for 1926-27, 1928, 15.) Isolated from cotton. Pseudomonas amaranti (sic) Smith. (U. S. Dept. Agr., Div. Veg. Phys. and Path. Bull., 28, 1901, 153; Bacterium amaranthi Smith, Bact. in Relation to Plant Dis., 3, 1914, 148; Phytomonas amaranthi Bergey et al., Manual, 1st ed., 1923, 186.) Isolated from diseased amaranthus. Growth in culture similar to Xantho- monas campestris and Xanthomonas hya- cinthi. Pseudomonas alutacea Migula. (Led- crgelber Bacillus, Tataroff, Die Dorpater Wasserbakterien, Inaug. Diss., Dorpat, * Prepared by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, July, 1943. FAMILY PSEUDOMOXADACEAE 179 1891, 61 ; Migula, Syst. d. Bakt., 2, 1900, 936.) Isolated from water. Pseudomonas graveolans Migula . ( Ba- cillus aquatilis graveolens Tataroff, Die Dorpater Wasserbakterien, Inaug. Diss., Dorpat, 1891, 48; Migula, Syst. d. Bakt., 2, 1900, 934.) Isolated from water. Not Pseudomonas graveolens Levine and Anderson (Jour. Bact., 23, 1932, 343) isolated from musty eggs, and by Olsen and Hammer (Iowa State Coll. Jour. Sci., 9, 1934, 125) from milk. Pseudomonas resinacea Migula. (Harzfarbener Bacillus, Tataroff, Die Dorpater Wasserbakterien, Inaug. Diss., Dorpat, 1891, 64; Migula, Syst. d. Bakt., 2, 1900, 935.) Isolated from water. Xanthomonas taraxaci Niederhauser. (Phytopath., 53, 1943, 961.) Patho- genic on Russian dandelion (Taraxacum kok-saghz) . Genus III. Methanomonas Orla-Jensen.* (Cent, f . Bakt., II Abt., 22, 1909, 311.) Cells monotrichous, capable of obtaining energy from oxidation of methane to CO2 and water. The type species is Methanomonas methanica (Sohngen) Orla-Jensen. 1. Methanomonas methanica (Sohn- gen) Orla-Jensen. (Bacillus niethanicus Sohngen, Cent. f. Bakt., II Abt., 15, 1906, 513; Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 311.) From methane. Short rods: 0.5 to 0.8 by 2.0 to 3.0 mi- crons, motile in young cultures by means of a single flagellum. In older cultures nearly spherical. Can be cultivated in an atmosphere composed of one part CH4 and two parts air on washed agar contain- ing the necessarj^ inorganic salts. The growth is membranous. At the end of two weeks, the organisms changed an atmosphere containing 225 ml. CH4 and 321 ml. O. to the follow- Clli 0 ml. , CO2 78 ml. O2 172 ml. In addition, 21 ml. CO2 was dissolved in the liquid. Habitat : Presumably widely distrib- uted in soil. Genus IV. Acetobacter Beijerinck.j (Proc. Kon. Akad. v. Wetenschapp., Amsterdam, 2, 1900, 495.) Acetobacter aceti first appeared (Krai's Sammlung v. Mikroorg., Prague, 1898, 4) as a synonym of Bacterium aceti Hansen. Beijerinck (loc. cit.) mentions Acetobacter aceti in a footnote of a later paper. The genus name Acetobacter was accepted by Fuhrmann (Beiheft Bot. Centralbl., Orig., 19, 1905, 8) and others. From Latin, acetum, vinegar; baclrum, rod. Synonyms: ?Ulvina Kiitzing, Algae aquae dulcis, etc., 11th decade, 1837; Myco- derma Thompson, Ann. d. Chem. u. Pharmacie, 83, 1852, 89 ; ? Umbina Naegeli, Bericht liber die Verhandlingen der bot. Section der 33 Versammlung deutscher Xatur- forscher. und Arzter. Bot. Ztg., 1857, 760; Bacterium Lanzi, N. Giorn. bot. ital., 1876, 257; Torula Saccardo, Atti Soc. Ven. Trent., 5, 1878, 315; Bacteriopsis (in part)Trevi- san, Atti Accad. Fisio-Medico-Statistica Milano, Ser. 4, 3, 1885, 103; Micrococcus Maggi, Jour. Microg., 10, ISSQ; Bacillus Schroeter, Kryptogamen Flora von Schlesien, 3, 1, 1886, 161; Termobacterium Zeidler, Cent. f. Bakt., II Abt., 2, 1896, 739; Acetobac- * Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, December, 1922. t Revised by Dr. C. D. Kelly, McGill Univ., Montreal, P. Q., Canada, July, 1938; further revision by Dr. Reese H. Vaughn, Univ. of California, Berkeley, California June, 1943. 180 MANUAL OF DETERMINATIVE BACTERIOLOGY terium Ludwig, in abstract of Hoyer's Inaug. Diss., Cent. f. Bakt., II Abt., 4, 1898, 867; Acetimonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 312. In addition, the sub-generic names E uacetobacter and Acetogluconobacier have been proposed by Asai, Jour. Agr. Soc. Japan, 11, 1935, 502. Tlie genus Gluconobacter and the sub-genera Eugluconobacter and Gluconoacetobacter Asai {loc. cit.) may be synonyms in whole or in part. Individual cells ellipsoidal to long and rod-shaped, occurring singly, in pairs, or in short or long chains. Motile with polar flagella, or non-motile. Involution forms may be spherical, elongated, filamentous, club-shaped, swollen, curved or even branched. Young cells Gram-negative; old cells often Gram-variable. Obligate aerobes; as a rule strongly catalase positive, sometimes weakly so. Oxidize various organic compounds to organic acids and other oxidation products which may undergo further oxidation. Common oxidation products include acetic acid from ethyl alcohol, gluconic and sometimes ketogluconic acid from glucose, dihydroxyacetone from gly- cerol, sorbose from sorbitol, etc. Nutritional requirements vary from simple to com- plex. Development generally best in yeast infusion or yeast autolysate media with added ethyl alcohol or other oxidizable substrate. Optimum temperature variable with the species. Widely distributed in nature where they are particularly abun- dant in plant materials undergoing alcoholic fermentation; of importance to man for their role in the completion of the carbon cycle and for the production of vinegar. The type species is Acetobacter aceti (Kiitzing) Beijerinck. Key to species of genus Acetobacter. I. Oxidize acetic acid to carbon dioxide and water. A. Capable of utilizing ammonium salts as a sole source of nitrogen (Hoyer's solution).* 1. Acetobacter aceti. B. Do not utilize ammonium salts as a sole source of nitrogen.* 1. Forms a thick, zoogloeal, cellulose membrane on the surface of liquid media. 2. Acetobacter xylinum. 2. Do not form a thick, zoogloeal membrane on the surface of liquid media. 3. Acetobacter rancens. 3a. Acetobacter pasteurianum. 3b. Acetobacter kuetzingianum . II. Do not oxidize acetic acid. A. Form pigments in glucose media. 1. Dark brown to blackish pigment. 4. Acetobacter melanogenum. 2. Pink to rose pigment. 5. Acetobacter roseinn. B. Do not form pigments. 1. Optimum temperature 30° to 35°C. 6. Acetobacter suboxydans. 2. Optimum temperature 20° to 25°C. 7. Acetobacter oxydans. * It is not known with certainty whether Acetobacter pasteurianum and Acetobacter kuetzingianum are capable of using inorganic nitrogen as a sole source of nitrogen for growth. However, since these two species are among those first described it is advisable to retain them for the present. See Acetobacter rancens Beijerinck. FAMILY PSEUDOMOXADACEAE 181 1. Acetobacter aceti (Kiitzing) Beijer- inck. (Ulvina aceti Kiitzing, Algae aquae dulcis etc., Uth decade, 1837; Mycoderma aceti Thompson, Ann. d. Chem. u. Pharmacie, 83, 1852, 89; Um- bina aceti Naegeli, Bericht iiber die Verhandlingen der bot. Section der 33 Versammlung deutscher Naturforscher und Arzter. Bot. Ztg., 1857, 760; Bac- terium aceti Lanzi, N. Giorn. bot. ital., 1876, 257; Torula aceti Saccardo, Atti Soc. Ven. Trent., 5, 1878, 315; Bacteriop- sis aceti Trevisan, Atti della Accademia Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 103; Micrococcus aceti Maggi, Jour. Microg., 10, 1886; Bacillus aceti Schroeter, Kryptogamen Flora von Schle- sien, 3, 1, 1886, 161; Bacillus aceticus Fliigge, Die Mikroorganismen, 1886, 313; Beijerinck, Krai's Sammlung v. Mikro- org., Prague, 1898, 7; Beijerinck, Proc. Kon. Akad. v. Wetensch., Amsterdam, 2, 1900, 495; Bacterium hansenianum Chester, Man. Determ. Bact., 1901, 126.) From Latin acetum, vinegar. Rods: 0.4 to 0.8 by 1.0 to 2.0 microns, occurring singly and in long chains, fre- quently showing large club-shaped forms. Stain yellow with iodine solution. Mo- tility variable. Motile cells possess a single polar flagellum (Vaughn, Jour. Bact., 46, 1943, 394). Forms large, shiny colonies on beer gelatin containing 10 per cent sucrose. Forms slimy pellicle on fluid media, or ring or turbidity without pellicle. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, sorbose, sucrose, maltose, lactose, raffinose, dex- trin, starch, glycogen, inulin, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, glycerol, erythritol, mannitol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Es- sigind., B, 1898, 147). Aerobic. Distinctive characters : Marked oxida- tive power causing rapid and complete oxidation of substrate as glucose or ethyl alcohol; ability to utilize inorganic nitro- gen salts as a sole source of nitrogen (Hoj^er, Inaug. Diss., Leiden, 1898, 43; Beijerinck, Cent. f. Bakt., II Abt., 4, 1898, 215) ; growth and oxidative activity in association with fermenting yeasts (Vaughn, Jour. Bact., 36, 1938, 360). Optimum temperature 30°C. Growth occurs between 10° and 42°C. Habitat: Vinegar; souring fruits, vege- tables and beverages. 2. Acetobacter xylinum (Brown) Hol- land. (Bacterium xylinum Brown, Jour. Chem. Soc, London, 49, 1886, 439; Holland, Jour. Bact., 5, 1920, 216; Ba- cillus xylinus Holland, ibid., 221.) From Gr. xijlinus, wooden (in reference to the cellulose in the membrane). Rods, about 2 microns long, occurring singly and in chains. The cells have a slimy envelope which gives the cellulose reaction. A film forms on the surface of liquids. This film becomes cartilagenous and falls to the bottom. This zoogloeal film forms on all liquid media in which growth occurs; the nature of the medium influ- ences the thickness of the film which may vary from 1 to 250 millimeters. X-ray pattern studies made by Khou- vine, Champetier and Sutra (Compt. rend. Acad. Sci. Paris, 194, 1932, 208) and by Barsha and Hibbert (Can. Jour. Research, 10, 1934, 170) have shown that the cellulose contained in the membranes formed by Acetobacter xylinum is identi- cal with cotton cellulose. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, maltose, lactose, raffinose, dextrin, starch, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, manni- tol and acetaldehyde (Henneberg, Die deutsch. Essigind., 2, 1898, 147). Aerobic. Distinctive character: The production of thick, leathery, zoogloeal cellulosic membranes on the surface of liquids. Optimum temperature 28°C. 182 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : Vinegar; souring fruits, vege- tables and beverages. 3. Acetobacter rancens Beijerinck. {Bacterium rancens Beijerinck, Cent. f. Bakt., II Abt., 4, 1898, 211; Beijerinck, Krai's Sammlung v. Microorg., Prague, 1898, 4.) From L. rancens, being rancid. Beijerinck {loc. cit.) in a footnote stated that "two of the many varieties of B. rancens have been described by Hen- neberg under the names of B. oxydans and B. acetosum. Hansen erroneously called this species B. aceti as did Brown. Neither Hansen nor Brown knew B. aceii Pasteur." No further morphologi- cal description is given. The following description is taken in part from a study of a culture of Aceto- bacter rancens received from Kluyver (Vaughn). Rods with the usual morphological ap- pearance of cultures of acetic acid bacteria. Gram-negative. Motility variable. Motile cells possess a single polar flagellum (Vaughn, Jour. Bact., 46, 1943, 394). Involution forms com- monly appear as filaments and enlarged cells. Wort agar slant: Growth abundant, butyrous, pale-buff in color in one week. Yeast infusion, glucose, calcium carbonate slant : Growth abundant, bu- tyrous and cream-colored in one week. With petri dish cultures well isolated colonies are large, smooth and butyrous on either medium. Broth cultures containing peptone or yeast infusion form a mucilaginous, slimj^ pellicle. Beijerinck {loc. cit.) called this polysaccharide pellicle, cellulose-like and intimated that the mucilaginous material in the pellicle was somewhat different from that produced by Aceto- bacter xylinum. The pellicle material stained blue when treated with iodine and hydroiodic acid. Acid from glucose, ethyl alcohol, propyl alcohol, butyl alcohol, glycol, adonitol, mannitol and sorbitol. No acid from numerous other compounds tested. Distinctive character : Production of a thin, mucilaginous, slimy, polysaccharide membrane on the surface of liquids as compared with the tMck, true cellulose membrane of Acetobacter xylinum grown under the same conditions. Beijerinck {loc. cit.) reported the production of a cellulose-like membrane with some cul- tures of Acetobacter rancens. Source : Isolated from shavings in the quick vinegar process. Habitat : Found in fermented grain mash, malt beverages, mother of vinegar. Beijerinck (Cent. f. Bakt., II Abt., 4, 1898, 211) thought that the next two species were hardly more than varieties of Acetobacter rancens. 3a. Acetobacter pasteurianum (Han- sen) Beijerinck. {Mycoderma ■pasteuri- anum Hansen, Compt. rend. d. Trav. d. Lab. d. Carlsberg, 1, 1879, 96; Bacteriuvi pasteurianum Zopf, Die Spaltpilze, 2 Aufl., 1884, 49; Beijerinck, Krai's Samm- lung V. Microorg., Prague, 1898, 7.) Named for Pasteur, the French chemist and bacteriologist. Rods: 0.4 to 0.8 by 1.0 micron, occurring singly and in chains, at times showing thick, club-shaped forms. Mo- tility variable. Motile cells possess a single polar flagellum (Vaughn, Jour. Bact., 46, 1943, 394). Stains blue with iodine. Wort gelatin colonies: Small, circular, entire, gray, slimy. Forms a dry, wrinkled folded pellicle on double beer with one per cent alcohol. Meat infusion gelatin: Widespread, later rosette form, toothed. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, sorbose, sucrose, maltose, lactose, raffinose, dextrin, starch, glycogen, inulin, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amj'l alcohol, glycerol, erythritol, mannitol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind., 2, 1898, 147). Aerobic. FAMILY PSEUDOMONADACEAE 183 Optimum temperature 30°C. Growth occurs between 5° and 42°C. Habitat : Vinegar; beer and beer wort. 3b. Acetobacter kuetzingianum (Han- sen) Bergey et al. (Bacterium kuetzingi- anum Hansen, Compt. rend. d. Trav. d. Lab. d. Carlsberg, 3, 1894. 191; Bergey et al., Manual, 1st cd., 1923, 35.) Xamed for Kuetzing, the German botanist. Short, thick rods, occurring singly. Rarely forming chains of notable length. Capsule stained blue with iodine and witli potassium iodide. Xon-motile. Double beer gelatin colonies : Small, entire, with vermiform surface. Wort gelatin colonies: Small, entire, with surface free of wrinkles. Double beer: Forms a rather thick, folded pellicle. Distinguished from A ce- tobacter aceti in showing heavier growth above the surface of the media. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, sorbose, sucrose, maltose, lactose, raffinose, dex- trin, starch, glycogen, inulin, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, glycerol, erythritol, mannitol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind., £, 1898, 147). Aerobic. Optimum temperature 34°C, maximum 42°C, minimum 6 to 7°C. Habitat : Beer. Found in double beer. gelatin becomes insoluble in boiling water and in trypsin solution. Beer- or wort-gelatin plates: Charac- teristic dark brown, wide-spreading, diffuse areas. Tap water-agar-glucose-peptone-po- tassium phosphate-iron citrate-chalk medium: In 24 hours at 30°C, black, spreading, diffuse areas. Utilizes peptone as a source of nitrogen. Produces the pigment from peptone only if maltose or glucose is present as a source of carbon. When grown in glu- cose-peptone broth with CaCOs at 25° to 30°C, black pigment is produced after several weeks, and the carbonate is changed to calcium gluconate. Pigment : The pigment causing the brown coloration is an aromatic substance which is blackened by iron salts. Re- duces alkaline solutions of silver and mercury, blackening them. Oxidizes mannitol and sorbitol to fruc- tose and sorbose. Does not attack sucrose and fructose. ]\Iuch gluconic acid is produced. Acid from glucose and mal- tose. Acetic acid produced from alcohol. Distinctive character: The formation of dark brown to black pigment in media containing a suitable substrate; par- ticularly glucose. Source : Isolated from beer. Habitat: Causes light-colored beer to become darker brown. It is a very strong beer-vinegar bacterium. Also found in souring fruits. 4. Acetobacter melanogenum Bei- jerinck. (Cent. f. Bakt., II Abt., 29, 1911, 175.) From Greek melas (mela7i), black; -genes, producing. Rods : Xon-motile or motile. Motile cells possess a single polar flagellum (Vaughn. Jour. Bact., 46, 1943, 394). Gelatin: Apparent liquefaction prob- ably caused by acid, not an enzyme. When hold on artificial media for some time, the power of liquefying gelatin is lost, probably due to a slower production of acid. Deep brown pigment produced ; 5. Acetobacter roseum Vaughn. {Bacterium hoshigaki var. rosea Taka- hashi and Asai, Cent. f. Bakt., II Abt., 82, 1930, 390; Acetobacter fioshigaki Bergey et al., :Manual, 4th ed., 1934, 39; Vaughn, Wallerstein Lab. Communica- tions, o, Xo. 14, 1942, 20.) From Latin, rasa, rose. Rods : 0.7 to 0.9 by 1.5 to 1.8 microns, generally occurring singly, at most in pairs, often in chains. Xon-motile. Pellicle on fluid media yields no starch or cellulose reaction. 184 MANUAL OF DETERMINATIVE BACTERIOLOGY Koji (a mixture of rice and mold spores used to start fermentation of Japanese bread and sak6) extract agar colonies : Small, granular, circular, glistening, umbonate, becoming brownish. Wort agar colonies : Circular, milky- white, becoming brownish in center and yellowish at periphery. Glucose sake agar: Circular, milky- white, granular, umbonate, entire. Hoshigaki (dried persimmons) extract agar: Circular, milky -white, granular, becoming yellowish-brown in the center and grayish-white at the periphery. Koji extract agar streak : Grayish- white, glistening with ciliate margin, becoming purple brown to brown. Koji extract: Turbid with thin film, ascending on wall of tube. Bouillon: Turbid with ring formation. Yeast infusion glucose agar : Colonies similar to those on wort agar. Yeast infusion glucose broth : Turbid with thin, ascending film. Red color produced on sake wort agar and all media containing calcium carbonate. Acid from glucose, fructose, galactose, arabinose, glycerol, mannitol, ethyl and propyl alcohol. No acid from maltose, sucrose, lactose, raflinose, dextrin, starch, inulin, sorbitol, glycogen, isodulcitol and methyl alcohol. Forms gluconic acid from glucose. Aerobic. Optimum temperatures 30° to 35°C; maximum 40° to 41°C; minimum 10° to 15°C. Thermal death point 50°C for 5 minutes. Distinctive character : The formation of a rose to red pigment in suitable media ; particularly those containing glucose and calcium carbonate. Source : Isolated from fermenting mash of dried persimmons (hoshigaki), and souring figs and dates. Note: Vaughn, Wallerstein Lab. Com- munications, 5, No. 14, 1942, 20, has proposed the name Acetohacter roseiun to replace the name Acetohacter hoshigaki. As originally described, this organism was given the name Bacterium hoshigaki var. rosea by Takahashi and Asai (loc. cit.) without the authors having first named and described the species Bac- terium hoshigaki. The Japanese word "hoshigaki" has been used in a confus- ing manner viz. Takahashi and Asai, loc. cit. {Bacterium industrium var. hoshigaki) and Takahashi and Asai, Jour. Agr. Chem. Soc. Japan, 9, 1933, 351 and Cent. f. Bakt., II Abt., 87, 1933, 385 {Bacteriuin hoshigaki var. glucuronicum I, II and III). None of these Japanese names are in the form of true binomials. 6. Acetobacter suboxydans Kluyver and de Leeuw. (Paper read at the con- vention of the Dutch Society of Microbiol- ogy, Utrecht, December, 1923, see Tijd- schrift V. Vergelijkende Geneeskunde, 10, Afl. 2-3, 1924.) From L. sub, under, less; Gr. oxys, sharp, acid; dans, giving, i.e. less acid giving; less oxidizing. Short rods : Occurring singly or in chains. Non-motile. Morphologically like Acetobacter rancens. Forms very thin, hardly visible pellicle on fluid media. Wort agar colonies: Very small, circu- lar, slightly yellow. Acid from ethyl alcohol, propyl alco- hol, glycol, glucose, glycerol and sorbitol. Optimum temperature 30°C. Distinctive character: Partial oxida- tion of substrates as indicated by the formation of calcium 5-keto gluconate crystals on the surface of agar slants con- taining glucose and calcium carbonate. Source : Isolated from spoiled beer. Habitat : Beer. 7. Acetobacter oxydans (Henneberg) Bergey et al. (Bacteritmi oxydans Hen- neberg, Cent. f. Bakt., II Abt., 3, 1897, 223; Bacillus oxydans Migula, Syst. d. Bakt., 2, 1900, 800; Bergey et al.. Manual, 1st ed., 1923, 36.) From Gr. oxys, sharp, acid; dans, giving. Rods: 0.8 to 1.2 by 2.4 to 2.7 microns, occurring singly and in chains. Motile FAMILY PSEUDOMONADACEAE 185 cells possess a single polar flagellum (Vaughn, Jour. Bact., ^6, 1943, 394). The chains show bud-like swellings. Gelatin colonies : Circular, becoming ir- regular in shape with peculiar ramifica- tions. Acid from arabinose, fructose, glucose, galactose, sucrose, maltose, raffinose, dextrin, ethyl alcohol, propyl alcohol, erythritol, mannitol, glycol and glycerol. No acid from sorbose, lactose, starch, glycogen, inulin, methyl alcohol, isopro- pyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind.,;2, 1898, 147). Aerobic. Optimum temperature 18° to 21°C. Distinctive characters: Low optimum temperature for growth and oxidation of substrates; and the ability to oxidize a large number of substrates. Habitat: Beer. Appendix: The following species have been described, but until more compara- tive studies have been made, no change in nomenclature is recommended or advisable. 1. Acetobacter zeidleri Beijerinck. (Termobaderium aceti Zeidler, Cent. f. Bakt., II Abt., 2, 1896, 739; Bacterium zeidleri Beijerinck, Acetobacter zeidleri Beijerinck, Krai's Sammlung v. Mikro- org., Prague, 1898, 7; Bacillus zeidleri Migula, Syst. d. Bakt., 2, 1900, 801; Acetobacter lindneri Bergey et al., Man- ual, 1st ed., 1923, 36.) Named for A. Zeidler, who first isolated this species. Rods, occurring singly and in chains, showing large sausage-shaped involution forms. Motile with a single polar flagel- lum (Zeidler, Cent. f. Bakt., II Abt., 4, 1898, 669). Wort gelatin: Small, circular, slightly granular, yellowish-brown, entire colonies. No liquefaction. Dirty, yellowish-brown pellicle on liquid media. Wort gelatin slant : Strongly glistening, transparent, whitish in center, smooth, very weakly liquefied. Potato: Very scant growth. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, maltose, lactose, rafhnose, dextrin, glycogen methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, glycerol, mannitol and acetaldehyde (Henneberg, Die deutsch. Essigind., 2, 1898, 147). Aerobic, facultative. Optimum temperature 25°C. Habitat: Beer wort. 2. Acetobacter acetosum (Henneberg) Bergey et al. {Bacterium acetosutn Hen- neberg, Cent. f. Bakt., II Abt., 3, 1897, 223; Bergey et al.. Manual, 1st ed., 1923, 36.) From Latin, acelum, vinegar. Rods : 0.4 to 0.8 by 1 .0 micron, occurring singly and in chains. Non-motile. Stains yellow with iodine. On beer, yeast water and glucose solu- tions a firm, coherent, uniform, smooth, white film that becomes folded (Henne- berg, Garungsbakt., 2, 1926, 201). Acid from glucose, galactose, ethyl alcohol, and propyl alcohol. No acid from arabinose, fructose, sorbose, sucrose, maltose, lactose, raffinose, dextrin, starch, glycogen, inulin, methyl alcohol, isopropyl alcohol, butyl alcohol, iso- butyl alcohol, amyl alcohol, glycerol, erythritol, mannitol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind., 2, 1898, 147). Optimum temperature 28°C, maximum 36°C, minimum 8°C (Henneberg, Cent, f. Bakt., II Abt., 4, 1898, 14). Habitat : Beer. 3. Acetobacter ascendens (Henne- berg) Bergey et al. {Bacterium ascen- dens Henneberg, Zeitschr. f. deutsche Essigind., Berlin, No. 19 to 23, 1898, 145; also see Cent. f. Bakt., II Abt., 4, 1898, 933; Bergey et al., Manual, 1st ed., 1923, 37.) From Latin, ascendo, pp. ascen- dens, ascending. Rods, occurring singly, rarely in chains. 186 MANUAL OF DETERMINATIVE BACTERIOLOGY Non-motile. Do not give the cellulose reaction with iodine solution. Glucose gelatin colonies : Dry, white, with white area surrounding the colony. Fluid cultures have a tough pellicle rising on the wall of the flask. Acid from ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, glucose, galactose, sucrose, maltose, lactose, raffinose, dextrin, starch, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutyl alcohol, amyl alcohol, glycerin, mannitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind., 2, 1898, 147). Aerobic. Optimum temperature 31°C. Habitat: Isolated from vinegar and from red wine. 4. Acetobacter plicatum Fuhrmann. (Beihefte z. bot. Centralbl., Orig., 19, 1905, 8.) Description given in Cent. f. Bakt., II Abt., 15, 1906, 377. From plicatus, folded. Rods : 0.55 to 0.7 by 0.75 to 0.9 microns when grown on agar at 28° to 30°C. Young streak cultures 0.4 to 0.6 by 1.4 to 1.6 microns with homogeneous staining when grown on beef-extract-gelatin at 22°C. 0.5 by 1.5 to 1.7 microns with uneven staining (polar) when grown on wine gelatin. At about 40°C the organ- isms form swollen and greatly elongated forms. Non-motile. Agar slant : Pale yellowish, translucent growth. Alcohol-free beer with glucose and sucrose: Turbid with thick pellicles. Potato : Growth limited. Ferments alcohol to form acetic acid. Optimum temperature 28° to 30°C. Habitat: Wine. 5. Acetobacter acetigentmi (Henne- berg) Bergey et al. {Bacterium aceti- genum Henneberg, Cent. f. Bakt., II Abt., 4, 1898, 14; Bacillus aceligenuni Migula, Syst. d. Bakt., 2, 1900, 801; Bergey et al., Manual, 1st ed., 1923, 35.) From Latin, producing vinegar. Rods, occurring singly and in pairs. 0.8 to 1.2 by 1.2 to 1.4 microns. Motile. Cells give a cellulose reaction with H0SO4 and iodine. Glucose gelatin colonies : Raised, gray- ish, slimy. Fluid cultures show a tough, slimy pellicle. Acid from glucose, ethyl alcohol, propyl alcohol and glycol. No acid from arabinose, fructose, galactose, sor- bose, sucrose, maltose, lactose, raffinose, dextrin, starch, glycogen, inulin, methyl alcohol, isopropyl alcohol, butyl alcohol, isobutjd alcohol, amyl alcohol, glycerol, erythritol, mannitol, dulcitol and acetal- dehyde (Henneberg, Die deutsch. Essig- ind.,;2, 1898, 147).' Aerobic. Optimum temperature 33°C. Thermal death point 43° to 45°C for 5 minutes. Habitat : Vinegar. 6. Acetobacter industrium (Henne- berg) Bergey et al. {Bacterium indus- trium Henneberg, Zeitschr. f. deutsche Essigindustrie, Berlin, 1898; Cent. f. Bakt., II Abt., 4, 1898, 933; Bacillus industrius Migula, Syst. d. Bakt., 2, 1900, 801; Bergey et al.. Manual, 1st ed., 1923, 36.) From Latin industrius, diligent. Rods: 0.3 to 0.8 by 2.4 to 20 microns, occurring singly and in chains. No distinct color produced with iodine. Motile. Forms pellicle on fluid culture media. Acid from arabinose, fructose, glucose, galactose, sucrose, maltose, lactose, raffi- nose, starch, dextrin, ethyl alcohol, propyl alcohol, glycol, glycerol and mannitol. No acid from isopropyl alco- hol, butjd alcohol, isobutyl alcohol, amyl alcohol and acetaldehyde (Henneberg, Die deutsch. Essigind., 2, 1898, 147). Aerobic. Optimum temperature 23°C. Maxi mum 35°C. Minimum 8°C. Habitat : Beer wort. FA&ULY PSEUDOMONADACEAE 187 7. Bacterium schuezenbachii Henne- berg. (Die deutsche Essigind., No. 11-18, 1906; also Cent. f. Bakt., II Abt., 17, 1906, 790.) Named for Schiizenbach, the inventor of the German quick vinegar process. Rods: 0.3 to 0.4 bj^ 1.0 to 3.6 microns, occurring singly, in pairs and chains. The cells are round, oval or elongated, not infrequently sickle-shaped or irregularly bent with rounded or pointed ends. Not stained with iodine. Non-motile. Wort gelatin colonies : Round, shin}', transparent with yellowish-brown cen- ters. A non-coherent fihn produced on the surface of liquid media. Acid from arabinose, fructose, glucose, galactose, maltose, lactose, dextrin, ethyl alcohol, propyl alcohol, glycerol and erythritol. Small amount of acid from sucrose and raffinose. No acid from mannitol (Henneberg, Handbuch d. Gar- ungsbakt., 2, 1925, 239). Temperature relations : Optimum 25° to 27.5^0. Scant growth at 34° to 35"C and 13° to 15°C. Xo growth at 37° and 7.5°C. Source: Isolated from vinegar in the quick vinegar process. Habitat: Produces acetic acid in quick vinegar process. 8. Bacterium xylinoides Henneberg. (Die deutsche Essigind., No. 11 to 18, 1906; also Cent. f. Bakt., II Abt., 17, 1906, 794.) From Greek, woody. Rods: 0.5 to 0.8 microns (round cells) and 0.5 to 1.2 microns (long forms), oc- curring singly, in pairs or chains, cells round and as short and long rods. The thick membrane like that produced by Acetohactcr xylinum gives the reaction for cellulose with iodine and sulfuric acid, but the thin membrane does not. Wort gelatin : Colonies are produced like drops of water, often with light brown kernels in the center. Wort gelatin streak : Growth transpar- ent at first, later whitish. Three types of membrane on fluid media are formed by this species and all three may even be found on a culture at one time. A thin, firm, coherent mem- brane like that of Bacterium orleanense may be formed or one that is voluminous, scumlike (like coagulated egg-white), coherent, somewhat slimj'^ and finally thick like that of Bacterium xylinum. Also a membrane may be formed that is intermediate in type. Acid from arabinose, glucose, galac- tose, sucrose, maltose, ethyl alcohol, propyl alcohol, glycerol and erythritol. Small amount of acid from fructose and mannitol. Temperature relations : Optimum 28°C. Slight growth at 14°C. No growth at 6°C. Source : Isolated from wine vinegar from the Rhine and other sections. Habitat : Found in vinegar made by the Orleans method. 9. Bacterium orleanense Henneberg. (Die deutsche Essigind., No. 11-18, 1906; also Cent. f. Bakt., II Abt., 17, 1906,792.) Latinized, of Orleans. Rods: 0.4 to 0.5 by 1.2 to 2.1 microns, occurring singly or in chains. The cells are round, elongated or as involution forms, with straight or curved cells appearing. Not stained with iodine. Non-motile. Wort gelatin : Colonies irregular in form, whitish in color, about 1 mm. in diameter in 2 days. Wort gelatin streak: Growth often slimy, transparent, licjuid mass with yellowish-brown sediment. Wort agar streak: Some strains form isolated, moist, slimy, transparent colo- nies and on the water of condensation isolated whitish colonies are formed. Other strains form a coherent, trans- parent coating with a light brown pre- cipitate later and individual, distinct, round colonies of the same color. Characteristic of this species is the firm coherent film on the surface of liquid media. 188 MANUAL OF DETERMINATIVE BACTERIOLOGY Acid from arabinose, glucose, galac- tose, maltose, lactose, raffinose, dextrin, ethyl alcohol, propyl alcohol, glycerol, erythritol and mannitol. Small amount of acid from fructose and sucrose (Hen- neberg, Handbuch d. Garungsbakt., 2, 1926,239). Temperature relations : Optimum 20° to 30°C. Slight growth at 35° to 36°C and 14° to 15°C. No growth at 39° and at 7° to 8°C. Source : Isolated from vinegar in the quick vinegar process. Habitat: Can be used both in the quick or German process and the Orleans method of making vinegar. 10. Bacterium vini acetati Henneberg. (Die deutsche Essigind., No. 11-18, 1906; also Cent. f. Bakt., II Abt., 17, 1906, 797.) From Latin vinum, wine and acetum, vinegar. Rods: 0.3 to 0.8 by 0.8 to 2.0 microns, occurring singly, in pairs and sometimes as short chains of three; cell round, oval or slightly elongated, and rarely moder- ately long forms. Streptococcus-like cells are found on older agar cultures and spindle forms in beer gelatin with 10 per cent sucrose. Wort gelatin : Round, moist, shiny, transparent colonies with whitish sedi- ment in the center. The film on liquid media is not strongly coherent and the liquid is cloudy. Acid from arabinose, fructose, glucose, galactose, sucrose, maltose, raffinose, dextrin, ethyl alcohol, propyl alcohol, glycerol and erythritol. No acid from lactose (Henneberg, Handbuch d. Gar- ungsbakt., .?, 1925, 239). Optimum temperature 28° to 33°C. Source: Wine vinegar. Habitat : Found in vinegar made by the Orleans method for wine vinegar. 11. Bacterium curvum Henneberg. (Die deutsche Essigind., No. 11-18, 1906; also Cent. f. Bakt., II Abt., 17, 1906, 791.) From Latin, curvus, bent. Rods: 0.4 to 0.5 by 2.0 to 2.4 microns, occurring singly or in pairs, cells usually oval or elongated, not infrequently sickle- shaped, with rounded or pointed ends. Not stained with iodine solution. Non- motile. Wort gelatin : Transparent, round colo- nies with raised center and edge, fre- quently whitish and dry. A non-coherent scanty pellicle is formed on the surface of liquid media which sinks readily and the liquid is quite turbid. Forms round white islands on the surface of wort with 3 per cent alcohol. In old cultures on beer are to be found numerous smooth light brown raised colonies about 1 mm in diameter on the uniform transparent base of the surface membrane. Acid from arabinose, glucose, raffinose, dextrin, ethyl alcohol, propyl alcohol, glycerol and erythritol. Small amount of acid from fructose, galactose and mannitol. No acid from sucrose, maltose and lactose (Henneberg, Handbuch d. Garungsbakt., 2, 1925, 239). Temperature relations : Optimum 25° to 30°C. Scant growth at 16° to 17°C. No growth at 7° to 8°C. Growth at 35°C. No growth at 39°C. Source : Isolated from vinegar in the quick vinegar process. Habitat : Produces acetic acid in the quick vinegar process. 12. Acetobacter viscosum Shimwell. {Bacterium aceti viscosum Day and Baker, Cent. f. Bakt., II Abt., 36, 1913, 433; Bacillus aceti viscosum Day and Baker, ibid., 437; Also see Baker, Day and Hulton, Jour. Inst. Brewing, — , 1912, 651; Shimwell, Jour. Inst. Brewing, 42 (N. S. 32), 1936, 586.) From Latin, viscous or slimy. Rods: 0.4 by 1.2 microns which produce ropiness in beer. No capsules observed. Non-motile as a rule. Weakly Gram- positive. Source: From ropy beer. FAMILY PSEUDOMONADACEAE 189 13. Acetobacter capsulatum Shimwell. (Jour. Inst. Brewing, 4^ (N. S. 32), 1936, 585.) From Latin, capsulated. Coccoid rods, 0.8 to 1.0 micron in malt extract media. 0.6 to 1.5 microns in other media. Produce ropiness in beer. Cap- sulated. Motile. Gram-negative. Source: From ropy beer. 14. Acetobacter gluconicum (Her- mann).* (Bacterium gluconicum Her- mann, Biochem. Zeit., 192, 1928, 198; also see Hermann, Biochem. Zeit., 205, 1929, 297 and Hermann and Neuschul, Biochem. Zeit., 233, 1931, 129.) It is unfortunate that an organism so well described must be placed with other species of uncertain standing. How- ever, this organism is so closely related to the other organisms described in the literature that further study is nec- essary. Source: From kombucha, a mixture of fungi and bacteria from tea infusions. 15. Acetobacter Tosic and Walker. 48, 1942, 82.) turbidans Cosbie, (Jour. Inst. Brewing, This beer vinegar bacterium is char- acterized by the production of intense turbidity in beer and ale. The descrip- tion given does not, at present, warrant recognition of the organism as a new species. Source: From beer. 16. Bacterium dihydroxyacetonicum Virtanen and Biirlund. (Biochem. Zeit., 169, 1926, 170.) There is no adequate description of this bacterium, and it is doubtful whether it can be properly evaluated since various species of Acetobacter also possess the ability to produce dihydroxy- acetone from glycerol. Consideration of this as a nomen nudem was indicated by Virtanen to ^'aughn in a personal communication in 1938. Source: From beet juice. 17. Acetobacter peroxydans Visser 't Hooft. (Inaug. Diss., Delft, 1925, 98.) The exact taxonomic position of this bacterium will not be clear until further comparative studies have been made. Source: From hj'drogen peroxide solu- tions. Genus V. Protamiaobacter den Dooren de Jong.] (Bijdrage tot de kennis van het mineralisatieproces. Thesis, Rotterdam, 1926, 159.) From AI. L., protamine and Latin, bactrum, rod. Cells motile or non-motile. Capable of dissimilating alkylamins. Pigmentation frequent. Soil or water forms. The type species is Protamitiobacter alboflavum den Dooren de Jong. Key to the species of genus Protaminobacter. I. Non-motile. Gelatin colonies light yellow to colorless. 1. Protaminobacter alboflavum. II. Motile. Gelatin colonies red. 2. Protaminobacter rubrum. 1. Protaminobacter alboflavum den Abt., 71, 1927, 218.) From Latin albus, Dooren de .Jong. (Thesis, Rotterdam, white; ^at^i/s, yellow. 1926, 159; also see Cent. f. Bakt., II Rods: Non-motile. Gram-negative. * It is uncertain at present who first used this combination. t Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, June, 1929; further revision by Prof. Robert S. Breed, New York State Experiment Station, Geneva New York, April, 1943. 190 MANUAL OF DETERMINATIVE BACTERIOLOGY Gelatin colonies: Circular, dry, light yellow or colorless. Gelatin stab: No liquefaction. Agar colonies: Circular, opaque, pig- ment bright red, yellow, light gray or colorless. Amine agar colonies : Circular, white to dark yellow. See table below for list of organic sub- stances utilized. 2. ProtaminobacterrubrumdenDooren de Jong. (Thesis, Rotterdam, 1926, 159; also see Cent. f. Bakt., II Abt., 71, 1927, 218.) From Latin, ruber, red. Rods : Motile with single polar flagel- lum (Weaver, Samuels and Sherago, Jour. Bact., S5, 1938, 59). Gram-nega- tive. Gelatin colonies : Circular, red, dry. Gelatin stab : No liquefaction. Table I. — Organic Substances Utilized as a Source of Carbon by Varieties of Protaminobacter alboflavum ORGANIC ACIDS Acetic Valerianic a-crotonic Undecyclic Lactic |8-oxybutyric Succinic Formic Glutaric Adipic Fumaric Malic Tartaric Citric /S-phenylpropi- onic Quinic + 0 + + + + 0 + 0 + + + + -t- -t- + 0 + 0 + -1- + + + 0 0 + -!- AMINO COxMPOUNDS a a-alanin a - aminocapronic acid Leucin Propionamid . . Capronamid. . . Uric acid Hippuric acid ALCOHOL Ethyl SUGAR Glucose a 0 7 a 0 0 + 4- + 0 + 0 + + 0 0 + 0 + 0 + 0 + 0 + 0 0 0 + 0 0 0 + + -h + + -t- + 0 Ethyl Diethyl Propyl Isopropyl.. . Dipropj'l. . . Tripropyl... Butyl Isobutyl Diisobutyl. Amyl Diamyl Ethanol Glucosamin Benzyl a & -y + + + -h + 0 -l- + + 0 + 0 + + 0 + 0 0 + 0 + + + 4- + + 0 + + + 0 + 0 + + -f + + + + 0 + Catalase is formed. Aerobic, facultative. Optimum temperature 30 °C. Habitat : Soil and water. Note : The author recognizes four varieties of this species which he dif- ferentiates on the basis of organic sub- stances attacked (see Table) and pig- ment produced. Variety a shows light yellow growth on gelatin, bright red on agar and yellow on amine agar. Variety ^ is light yellow on gelatin, yellow on agar and dark yellow on amine agar. Variety 7 is light yellow on gelatin, light gray on agar and yellow on amine agar. Variety 8 is colorless on gelatin and agar and white on amine agar. Agar colonies: Circular, red, opaque. Amine agar colonies : Circular, dark red. The following organic acids are at- tacked : Acetic, lactic, jS-oxybutyric, glycerinic, succinic, malonic, formic, methyl formic, glutaric, maleinic, fu- maric, malic, tartaric, citric and quinic. The following amino compounds are attacked: Sarcosin, betain, hippuric acid, asparagine, propionamid, caprona- mid, lactamid, succinamid, allantoin and uric acid. Glucose is fermented. Catalase is formed. Aerobic, facultative. Optimum temperature 30°C. Habitat : Soil and water. FAMILY PSEUDOMONADACEAE 191 Genns VI. Mycoplana Gray and Thornton * (Cent. f. Bakt., II Abt., 73, 192S, S2.) From Greek, mykes, fungus; plane, a wan- derer or traveller. Cells branching, especially in 3'oung cultures. Frequently banded when stained. Capable of using phenol and similar aromatic compounds as a sole source of energy. Grow well on standard culture media. Type species Mycoplana dimorpha Graj^ and Thornton. Key to the species of genus Mycoplana. I. Gelatin not liquefied. 1. Mycoplana dimorpha. II. Gelatin liquefied. 2. Mycoplana hxdlata. 1. Mycoplana dimorpha Gray and Thornton. (Cent. f. Bakt., II Abt., 73, 1928, 82.) From Greek, di, two; morphos, forms. ■ Short, curved and irregular rods, 0.5 to 0.7 by 1.25 to 4.5 microns, showing branching especially in young cultures. fMotile, with long polar flagella. Gram- negative. Gelatin colonies : Circular, buft', smooth, resinous, entire. Gelatin stab : Xo liquefaction. Growth filiform. Agar colonies: Circular, buff, convex, smooth, glistening, entire. Agar slant: Filiform, white, convex, glistening, entire. Broth: Turbid, with surface ring. Nitrites not produced from nitrates but gas evolved in fermentation tubes. Starch hydrolyzed. No acid from carbohydrate media. Attacks phenol. Aerobic. Optimum temperature below 30 °C. Source: One strain isolated from soil. Habitat: Probably in soil. 2. Mycoplana bullata Gray and Thorn- ton. (Cent. f. Bakt., II Abt., 73, 1928, 83.) From Latin, bullatus, furnished with a boss or knob. Rods, curved and irregular, branching, 0.8 to 1.0 by 2.25 to 4.5 microns. fMotile with polar flagella. Gram-negative. Gelatin colonies : Circular, buff, smooth, glistening, edge diffuse. Partially lique- fied. Gelatin stab: Saccate liquefaction. Agar colonies : Circular, white, convex, smooth, glistening, entire. Agar slant: Filiform, white, convex, smooth, glistening, entire. Broth: Turbid. Nitrites not produced from nitrates. Gas, presumabl}^ N, in fermentation tubes. Starch not hydrolyzed. No acid from carbohj'drate media. Attacks phenol. Aerobic. Optimum temperature below 30 °C. Source: Two strains isolated from soil. Habitat: Probably in soil. * Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, June, 1929. t The original statements regarding the flagellation of the.se species are contra- dictory. The first reads "Polar, peritrichous; the second "Polar or peritrichous". Drawings given usually indicate peritrichous rather than polar flagellation. Further study is needed before these species can be properly placed in relation to other known species. — Editors. 192 MANUAL OF DETERMINATIVE BACTERIOLOGY TRIBE II. SPIRILLEAE KLUYVER AND VAN NIEL. (Cent. f. Bakt., II Abt., H, 1936, 346.) More or less spirally curved cells. Key to the genera of tribe Spirilleae. I. Generally motile by means of a single polar flagellum. A. Short, bent rods occurring singly or united into spirals. Genus I. Vibrio, p. 192. B. Slightly curved rods of variable length. Strict anaerobes which reduce sulfates to hydrogen sulfide. Genus II. Desulfovibrio , p. 207. C. Cells oxidize cellulose forming oxj^cellulose. Growth on ordinary culture media is feeble. 1. Long, slightly curved rods with rounded ends. Genus III. Cellvibrio, p. 209. 2. Short, curved rods with pointed ends. Genus IV. Cellfalcicula, p. 211. II. Generally motile by means of a tuft of polar flagella. Cells of varying thick- ness, and length and pitch of spiral, forming either long curves or portions of a turn. A. Oxidize inorganic sulfur compounds. Cells contain free sulfur granules. Genus V. Thiospira, p. 212. B. Not as above. Genus VI. Spirillum, p. 212. Genus I. Vibrio Muller* (Miiller, Vermium terrestrium et fluviatilum, 1, 1773, 39; Pacinia Trevisan, Att^ d. Accad. Fisio-Medico-Statisticain Milano, Ser. 4, 3, 1885, 83; Microspira Schroeter, in Cohn, Kryptogamen-Flora von Schlesien, 3, 1, 1886, 168; Pseudospira De Toni and Trevisan, Sylloge Fungorum, 8, 1889, 1018; Photobacterium Beijerinck, Arch, neerl. d. sci. exactes, 23, 1889, 401; Liquidovibrio Orla-Jensen, Cent. f. Bakt. II Abt., 22, 1909, 333; Solidovibrio Orla-Jensen, ibid.; Dicrospira Enderlein, Sitzber. Ges. naturf. Freunde, Berlin, 1917, 313.) From Latin, vibro, vibrate. Cells short, curved, single or united into spirals. Motile by means of a single polar flagellum which is usually relatively short; rarely, two or three flagella in one tuft. They grow well and rapidly on the surface of standard culture media. Aerobic to anaerobic species. Mostly water forms, a few parasites. The type species is Vibrio comma (Schroeter) Winslow et al. Key to the species of genus Vibrio. I. Gelatin liquefied. A. Nitrites produced from nitrates. 1. Indole is formed. a. Milk not coagulated. 1. Vibrio comma. 2. Vibrio berolinensis. aa. Milk coagulated. 3. Vibrio metschnikovii. * Revised by Prof. D. H. Bergey, Philadelphia, Pennsylvania, April, 1937; partial revision by Capt. Wm. C. Haynes, Sn.C, Fort Bliss, Texas, July, 1943 and by Lt. Col. A. Parker Kitchens, University of Pennsylvania, Philadelphia, Penna., Decem- ber, 1943. FAMILY PSEUDOMONADACEAE 193 2. Indole not formed. a. Milk not coagulated. 4. Vibrio tyrogenus. 5. Vibrio xenopus. B. Nitrites not produced from nitrates. 1. Indole is formed. a. Milk coagulated, peptonized. 6. Vibrio piscium. 2. Indole not formed. a. Milk acid, coagulated. 7. Vibrio proteus. 8. Vibrio wolfii. 9. Vibrio sputigeniis. 10. Vibrio liquefaciens. aa. Milk not coagulated. b. Growth on potato thin, barely visible. 11. Vibrio strictus. bb. Xo growth on potato. 12. Vibrio aquatilis. aaa. Action on milk not reported. b. Acid from glucose. Attacks naphthalene. 13. Vibrio neocistes. bb. No acid from carbohydrates. Attacks naphthalene. 14. Vibrio cuneatus. bbb. No acid from carbohydrates. Liquefies agar. 15. Vibrio granii. II. Gelatin not liquefied. A. Nitrites produced from nitrates. 1. Acid and gas from glucose. 16. Vibrio leonardii. 2. Acid but not gas from glucose. Liquefies agar. 17. Vibrio agarliquefaciens. B. Nitrites not produced from nitrates. 1. Acid from glucose. IS. Vibrio cyclosiles. 2. No acid from carbohydrates. 19. Vibrio percolans. C. Nitrite production not reported. 1. Requires the addition of ammonium sulfate for growth. Ammonium sulfate agar liquefied. 20. Vibrio andoi. 2. Do not require ammonium sulfate for growth. a. Indole not formed. b. Microaerophilic, becoming aerobic. 21. Vibrio fetus. bb. Aerobic, facultative. 22. Vibrio pierantonii . 1. Vibrio comma (Schroetcr) Wins- 479; Spirillum cholerae asiaticae Zopf, low et al. (Kommabacillus, Koch, Die Spaltpilze, 3 Aufl., 1S85, 69; Paci/i/a Berliner klin. Wochenschr., 21, 1884, c/ioieraeas/aiicae Trevisan, Atti d. Accad. 194 MANUAL OF DETERMINATIVE BACTERIOLOGY Fisio-Med.-Statistica in Milano, Ser. 4, 3, 1885, 84; Microspira comma Schroeter, in Cohn, Kryptogamen Flora v. Schlesien, 3, 1, 1886, 168; Vibrio cholerae Neisser, Arch. f. Hyg., 19, 1893, 199; Vibrio cholerae asiaiicae Pfeiffer, in Fliigge, Die Mikroorganismen, 2, 1896, 527; Winslow et al., Jour. Bact., 5, 1920, 204; Bacillus cholerae Holland, Jour. Bact., 5, 1920, 217; Bacillus comma Holland, ibid., 218; Spirillum cholerae-asiaticae Holland, ibid., 225; Vibrio cholerae-asiaticae Hol- land, ibid., 226.) From Latin, comma. Slightly curved rods, 0.3 to 0.6 by 1.0 to 5.0 microns, occurring singly and in spiral chains. Cells may be long, thin and delicate or short and thick. May lose their curved form on artificial culti- vation. Motile, possessing a single polar flagellum. Gram-negative. Gelatin colonies: Smajl, yellowish- white. Gelatin stab : Rapid napiform lique- faction. Agar colonies : Circular, whitish-brown, moist, glistening, translucent, slightly raised, entire. Agar slant: Brownish-gray, moist, glistening. McConkey's medium: Good growth, colonies colorless when young, soon pinkish, medium becomes darker red. Broth: Slightly turbid, with fragile, wrinkled pellicle and flocculent precipi- tate. Peptone water: Characteristic rapid growth, chiefly at surface, where after 6 to 9 hours, a delicate membrane is formed; little turbidity, deposit appar- ently derived from pellicle (Topley and Wilson, Princip. Bact. and Immun., 2nd ed., 1936, 388). Readily isolated from the surface film of 0.1 per cent peptone water. Litmus milk : Alkaline at the top and slightly acid at bottom; generally not coagulated; peptonized; reduced. Potato : Dirty -white to yellowish, moist, glistening, spreading. Blood serum: Abundant growth, some- times slow liquefaction. Blood agar: The blood pigment is di- gested forming a greenish zone around colonies; a true soluble hemolysin is not formed (the El Tor vibrio also di- gests blood pigment but in addition pro- duces a soluble hemolysin. Otherwise it is said to be indistinguishable from the typical cholera vibrio) . Indole is formed. Nitrites produced from nitrates. Cholera-red reaction, which depends on production of indole and reduction of nitrates is positive. Hydrogen sulfide is formed. Acid but not gas from glucose, fruc- tose, galactose, maltose, sucrose and mannitol. Slowly from glycerol. Does not attack lactose, inulin or dulcitol. Group I of Heiberg (Classification of Vibrio cholerae and Cholera-like Vibrios. Copenhagen, 1935) ferments mannose and sucrose but not arabinose. Hydrolyzes starch actively in alkaline media. High alkali but low acid tolerance; optimum pH 7.6 to 8.0; for isolation on Dieudonne's medium pH 9.0 to 9.6. Aerobic, grows best in abundant oxy- gen; under strict anaerobiosis may fail to grow altogether. Optimum temperature 37 °C. Maxi- mum 42°C. Minimum 14°C. Source: From intestinal contents of cholera patients in Egypt and India. Habitat : Intestinal contents of cholera patients and carriers. The relationships existing among the cholerigenic and non-pathogenic water vibrios, although studied intensively, have not yet been completely defined. As a working scheme, based on somatic (O) and flagellar (H) antigen studies, Gardner and Vankatraman (Jour. Hyg., 35, 1935, 262-282) suggest the following FAMILY PSEUDOMONADACEAE 195 Cholera group of vibrios. (Biochemically similar. Common H antigen.) 0-sub-group I. Non-hemolytic (goat cells). Cholera vibrios. Types — original, variant and middle. Hemolytic (goat cells). El Tor vibrios. Types — original and variant (Pmiddle). Linton (Bact. Rev., 4, 1940, 275) has outlined a classification of the vibrios based upon their protein and poly- saccharide structures. Using chemical methods, it was found that one poly- saccharide and one protein was com- monly obtained from each strain of vibrio; when exceptions occurred, it was invariably noted that the strain was undergoing dissociation. Given a single protein and polj'saccharide in each vibrio, it was possible to divide the strains into six groups, which were numbered in the order of their discovery as shown in the table. A chemical grouping of the cholerigcnic and water vibrios. Group Protein Type I I II I III II IV II V II VI I Polysaccharide Type I II II I III III The strains of Groups I and II possess the same protein and different poly- saccharides. These are derived from cases of cholera and have the serological O sub-groups II, HI, IV, V, VI and individual races (mostly hemolytic). Paracholera, cholera-like, and some El Tor vibrios. (Types \Yithin sub-groups underlined.) and biochemical characteristics of O- Group I, Vibrio cholera. Group I strains are more common than those of Group II, which have, however, been isolated from epidemics with a high mortality. The phospholipid fraction is common to both types when isolated in the early part of an epidemic, but is not found in strains of other groups. The harmless water vibrios, which are so heteroge- neous serologically (Taylor and Ahuja, Indian Jour. Med. Res., 26, 1938, 8-32) form a single chemical group with a homogeneous structure. They fall into Group III, which differs in its protein structure from the authentic cholera vibrios, and resembles Group II in its polysaccharide. The vibrios of Group IV, which came from El Tor and from chronic vibrio carriers are believed on epidemiological grounds to be harmless, although serological methods have failed to distinguish them from cholerigcnic vibrios. Group V, which, like III and IV, contains protein II, consists, like Group IV, of strains from chronic vibrio carriers. Group VI strains are only rarely isolated in nature and representa- tives of this group are generally found among collections of old laboratory strains. They appear to be the result of polysaccharide variation from Group I 196 MANUAL OF DETERMINATIVE BACTERIOLOGY after long-continued growth on artificial media. 2. Vibrio berolinensis Neisser. (Arch, f. Hyg., 19, 1S93, 200; Microspira bero- linensis Migula, in Engler and Prantl, Die natiirl. Pflanzenfam., 1, la, 1895, 33.) From M. L., the genitive of Berolina, the Latin name for Berlin. Curved rods, somewhat smaller than Vibrio comma. Frequently occurring in pairs. Motile, possessing a polar flagellum. Pleomorphic. Gram-nega- tive. Gelatin colonies: Small, grayish, slightly granular, fragmented. Gelatin stab: Slow, napiform lique- faction. Agar slant: Grayish-yellow, moist, glistening. Broth: Turbid, with gray pellicle. Litmus milk: No coagulation, no acid. Potato: Brownish streak. Indole is formed. Nitrites produced from nitrates. Not pathogenic for mice, pigeons or guinea pigs. Aerobic, facultative. Optimum temperature 37 °C. Mini- mum above 10 °C. Maximum less than 60 °C. Source : Isolated from filtered Spree river water. 3. Vibrio metschnikovii Gamaleia. (Gamaleia, Ann. Inst. Pasteur, 2, 1888, 482; Pacinia meischmkoffi. Trevisan, I generi e le specie delle Battcriacee, 1889, 23; Spirillum metschnikovi Sternberg, Man. of Bact., 1893, 511; Vibrio nord- hafen Pfuhl, Ztschr. f. Hyg., 22, 1894, 234; Microspira metschnikoffii Migula, in Engler and Prantl, Die nattirl. Pfian- zenfam., 1, la, 1895, 33.) Named for Metschnikoff, Russian bacteriologist. Probable synonyms : Vibrio schuyl- kiliensis Abbott, Jour. Exp. Med., 1, 1896, 424 {Microspira schxylkiliensis Chester, Manual Determ. Bact., 1901, 334); Vibrio dannbicus Heidcr, Cent. f. Bakt., H, 1893, 341 {Microspira danubica Migula in Engler and Prantl, Die natiirl. Pflanzenfam., 1, la, 1895, 33; Spirillum danubicum Holland, Jour. Bact., 5, 1920, 225). Curved rods, somewhat shorter and thicker than Vibrio comma. Motile. Gram-negative. Gelatin colonies : Like those of Vibrio comma. Gelatin stab: Rapid, napiform lique- faction. Agar slant: Yellowish, plumose, moist, glistening. Broth: Turbid, with thin, white pel- licle. Litmus milk: Acid, coagulated (eighth day); not peptonized. Potato: Delicate, brownish growth. Indole is formed. Nitrites produced from nitrates. Pathogenic for pigeons, fowls, and guinea pigs. Aerobic, facultative. Optimum temperature 37°C. Maxi- mum less than 45°C. Source : Isolated from fowl dead of a choleraic disease. Habitat: The intestinal contents of chickens, pigeons and other animals suf- fering from a cholera-like disease. 4. Vibrio tyrogenus (Fliigge) Hol- land. (Kasespirillen, Deneke, Deutsch. med. Wochnschr., 11, 1885, 33; Spirillum tyrogenum Fliigge, Die Mikroorganismen, 2 Aufl.. 1886, 386; Pacinia denekei Trevi- san, I generi e le specie delle Batteriacee, 1889, 23; Microspira tyrogena Migula, in Engler and Prantl, Die natiirl. Pflan- zenfam., 1, la, 1895, 33; Holland, Jour. Bact., 5, 1920, 225; Vibrio denekei Ilauduroy et al., Diet. d. Bact. Path., 1937, 541.) From Greek tyros, cheese; genes, produced from. Curved rods, rather smaller and more slender than Vibrio comma, often very long, closely wound spirals. Motile, possessing a polar flagellum. Gram- negative. FAMILY PSEUDOMONADACEAE 197 Gelatin colonies : Small, gray, granular, entire. Gelatin stab: Rapid, saccate liquefac- tion. Agar slant: Yellowish-white, plumose, glistening. Broth: Turbid. Litmus milk : Not coagulated. Potato: No growth. Indole not formed. Slight production of nitrites from nitrates. Aerobic, facultative. Optimum temperature 30°C. Source: Isolated from cheese. 5. Vibrio xenopus Schrire and Green- field. (Trans. RoA^al Soc. So. Africa, 17, 1930, 309.) From Xenopus, a genus of African toads. Spiral forms, occurring singly and in pairs. Non-motile. Gram-negative. Gelatin stab : Slow, cratcriform liciue- faction. Agar colonies : Small, white, glistening, slim}% entire. Agar slant : Grayish-white, slimy, entire. Broth: Turbid with flocculent sedi- ment. Litmus milk: Unchanged. Potato: Not reported. Indole is not formed. Nitrites are produced slowly from nitrates. Blood serum is peptonized. Starch is not hydrolj'zed. Acid from glucose, fructose, maltose, glycerol and sorbitol. Aerobic, facultative. Optimum temperature 37°C. Source : Found in abscess of pectoral muscle of African toad. 6. Vibrio piscium David. (Cent. f. Bakt., I Abt., Orig., 102, 1927, 46.) From Latin pise is, fish. Curved rods : 0.3 to 0.5 by 2.0 microns. Motile with a single polar flagellum. Gram-negative. Gelatin colonies : Circular, granular, opaque. Gelatin stab : Napiform liquefaction. Agar colonies : Yellowish, circular, smooth, entire, iridescent. Agar slant : Light yellow, transparent streak. Broth: Slight turbidity, with thin pellicle. Litmus milk : Soft coagulum. Pep- tonized, alkaline. Potato: Brownish-red streak. Indole is formed. Nitrites not produced from nitrates. Hydrogen sulfide formed. No action in sugar media. Pathogenic for frogs. Aerobic, facultative. Optimum temperature 18° to 20°C. Habitat : Causes epidemic infection in fish. 7. Vibrio proteus Buchner. (Kom- mabacillus der cholera nostras, Finkler and Prior, Deutsche med. Wochenschr., 1884, 632; Buchner, Sitzungsber. d. Gesel. f. Morph. u. Physiol., Miinchen, Heft 1, 1885, 10; Pacinia finkleri Trev- isan, Atti d. Accad. Fi sio -Medico - Statistica in Milano, Ser. 4, 3, 1885, 84; Microspira finkleri Schroeter, in Cohn, Krytogamen-Flora von Schlesien, 3, 1, 1886, 168; Spirillum finkleri Crookshank, Man. of Bact., 3rd ed., 1890, 282; Micro- spira protea Chester, Manual of Deter- minative Bacteriology, 1901, 338; Vibrio finkleri Holland, Jour. Bact., 5, 1920, 225.) From Greek, Proteus, a marine deity who had the power of assuming any shape he chose. Curved rods : 0.4 to 0.6 by 2.4 microns, often pointed at both ends. Motile, possessing a polar flagellum. Gram- negative. Gelatin colonies: Small, gray, circular, granular, entire. Gelatin stab: Rapid, saccate lique- faction. Agar slant : Dirty grayish, plumose. Broth: Turbid, with fetid odor. 198 MANUAL OF DETERMINATIVE BACTERIOLOGY Litmus milk : Slightly acid ; coagulated ; peptonized. Potato : Grayish, slimy layer. Indole not formed. Nitriites not produced from nitrates. Aerobic, facultative. Optimum temperature 30 °C. Source: Isolated from feces of pa- tients suffering from cholera nostras. Habitat : Intestinal contents in cholera nostras and cholera infantum. 8. Vibrio wolfii (Migula) Bergey et al. {Bacillus choleroides Wolf, Mlinch. med. Wochenschr., 40, 1S93, 693; Microspira wolfii Migula, Syst. d. Bakt., 2, 1900, 1001; not Microspira choleroides Migula, loc. cit., 992; Bergey et al., Manual, 1st ed., 1923, 80.) Named for Wolf, who first isolated this organism. Curved rods and S-shaped forms. Motile. Gram-negative. Gelatin colonies: Small, grayish-white, spreading. Gelatin stab: Infundibuliform lique- faction. Agar slant: Gray, moist layer. Broth: Turbid, witli gray pellicle. Litmus milk: Acid; coagulated. Potato : Yellowish-white layer. Blood serum: Rapid liquefaction. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 37 °C. Source : Isolated from cervical secre- tions in chronic endometritis. Gelatin colonies: Small, circular, slightly granular, yellowish, becoming brownish. Gelatin: Crateriform liquefaction. Agar slant: Grayish-white, moist. Broth: Turbid, no pellicle formed. Litmus milk: Acid; coagulated. Potato : Thin, gray layer, spreading. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 37 °C. Source : Isolated from sputum. 10. Vibrio liquefaciens (Migula) Ber- gey et al. (Bonhoff, Arch. f. Hyg., 19, 1893, 248; Microspira liquefaciens Mi- gula, Syst. d. Bakt., 2, 1900, 990; Bergey et al.. Manual, 1st ed., 1923, 81.) From Latin, liquefacio, to make liquid. Comma and S-shaped rods. Motile. Gram-negative. Gelatin colonies: Circular, with ir- regular margin, surrounded by a rose- colored zone. Gelatin stab : Slow, napiform lique- faction. Agar slant: Smooth, grayish, plumose. Broth : Turbid, with heavy grayish pellicle. Litmus milk: Acid; coagulated. Potato : Moist, brownish layer. Indole is not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 37 °C. Habitat : Water. 9. Vibrio sputigenus (Migula) Bergey et al. (Vibrio aus Sputum, Brix, Hyg. Rundschau, 4, 1894, 913; Microspira sputigena Mignla, Syst. d. Bakt., f, 1900, 981; Bergey ct al.. Manual, 1st ed.. 1923, 80.) From Latin, spuo {sputus), spu- tum; -genes, produced from. Slightly curved rods, about the same size and form as Vibrio comma, occurring singly, occasionally three or four in a chain. Motile. Possessing a polar fiagel- lum. Gram-negative. 11. Vibrio strictus Kutscher. (Ztschr. f. Hyg., 19, 1895, 469.) From Latin stringo (strictus), constricted. Markedly curved rods, of about twice the size of Vibrio comma. Motile. Gram-negative. Gelatin colonies: Small, circular, yel- lowish, with serrate margin. Gelatin stab: Slow, napiform to sac- cate liquefaction. Agar slant : Growth plumose, moist. Broth: Turbid, with gray pellicle. FAMILY PSEUDOMONADACEAE 199 Litmus milk: Not coagulated. Potato: Thin, barely visible layer. Blood serum is slowlj' liquefied. Indole is not formed. Nitrites not produced from nitrates. Pathogenic for guinea pigs. Aerobic, facultative. Optimum temperature 37 °C. Habitat : Water. 12. Vibrio aquatilis Giinther. (Deut- sche med. Woehenschr., 1S92, 1124; M icrospira aquatilis Migula, System der Bakterien, 2, 1900, 993.) From Latin, aquaticus, living in water. Curved rods, like Vibrio comma. Motile, possessing a polar flagellum. Gram-negative. Gelatin colonies: Circular, brownish, finely granular, entire. Gelatin stab : Crateriform liquefaction. Agar slant : Moist, grayish, glistening. Broth: Slightly turbid. Litmus milk : Not coagulated. Potato: No growth. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30 °C. Habitat : Water. 13. Vibrio neocistes Gray and Thorn- ton. (Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 92.) From Greek neos, new and kiste bo.x or ark. Here used as the equivalent of Newark, the name of a city in England. Curved rods: 0.5 to 1.0 by 1.0 to 4.0 microns. Motile with one to three polar flagella. Gram stain not recorded. Gelatin colonies : Liquefied. Gelatin stab : Liquefied. Medium red- dened. Agar colonies : Circular or amoeboid, buff to brownish, convex, smooth, glis- tening, entire. Agar slant :Filiform, fluorescent, raised, smooth, glistening, undulate. Broth: Turbid. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid from glucose. Attacks naphthalene. Aerobic, facultative. Optimum temperature. Habitat: Soil. 14. Vibrio cuneatus Gray and Thorn- ton. (Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 92.) From Latin, cuneo (cuneatus) wedge. Curved rods : 1.0 by 1.0 to 3.0 microns, the cells tapering at one extremity. Motile with one to five polar flagella. Gram-negative. Gelatin colonies: Liquefied. Gelatin stab: Liquefied. Agar colonies : Circular to amoeboid, white to buff, fiat to convex, smooth, translucent, border entire. Agar slant: Filiform, whitish, smooth, glistening. Indole not recorded. Nitrites not produced from nitrates. Starch not hydrolyzed. No acid from carbohydrate media. Attacks naphthalene. Aerobic, facultative. Optimum temperature 30° to 35°C. Source : One strain isolated from soil from Rothamsted, England. Habitat: Soil. 15. Vibrio granii (Lundestad) Stanier. (Bacterium granii Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 330; Achromo- bacter granii Bergey et al., Manual, 3rd ed., 1930, 222; Stanier, Jour. Bact., 42, 1941, 538.) Named for Prof. H. H. Gran, who first detected agar-liquefying bac- teria. Rods: 0.6 to 0.8 by 1.4 to 2.4 microns, with rounded ends, occurring singly, in pairs, and at times in short chains. Motile. Polar flagellate (Stanier, loc. cit.). Gram-negative. Fish-gelatin colonies : Punctiform, black, glistening. Fish-gelatin stab : Slow, crateriform liquefaction. Sea-weed agar colonies: Circular, flat 200 MANUAL OF DETERMINATIVE BACTERIOLOGY opaque, glistening, white, slimy, entire. Agar is dissolved. Fish-agar slant: Flat, white, elevated, glistening, undulate. Liquefied. Broth: Turbid with grayish-white, slimy sediment. Indole not formed. Nitrites not produced from nitrates. Starch usually hydrolyzed. No action on sugars. Aerobic, facultative. Optimum temperature 20° to 25 °C. Source : Sea water of Norwegian Coast. Habitat: Presumably sea water and on sea weeds. 16. Vibrio leonardii Metalnikov and Chorine. (Ann. d. I'Inst. Pasteur, ^2, 1928, 1647.) Named for Leonard. Curved rods with rounded ends, 0.5 to 1.0 by 2.0 to 3.0 microns. Motile with 1 to 3 polar flagella. Gram-negative. Gelatin stab : No liquefaction. Agar colonies: Small, transparent, cir- cular, having a characteristic odor. Broth: Turbid, with thin pellicle. Litmus milk: No coagulation, acid, with reduction of litmus. Potato: Slight, colorless growth. Indole not formed. Nitrites produced from nitrates. Blood serum not liquefied. Hydrogen sulfide formed. Acid and gas from glucose, fructose, galactose, lactose, sucrose and mannitol. No acid or gas from maltose or glycerol. Aerobic, facultative. Optimum temperature 30°C. Habitat : Highly pathogenic for insects as Galleria mellonella L. (bee moth), and Pyrausta nubialis Hiibn. (European corn borer). 17. Vibrio agarliquefaciens (Gray and Chalmers) Bergey et al. (Microspira agar-liquefaciens Gray and Chalmers, Ann. Appl. Biol., 11, 1924, 325; Manual, 4th ed., 1934, 119.) From Latin, lique- facio, liquefying; Malay, agar, a jelly from seaweeds. Short curved rods, usually c-shaped, with occasional s-shaped and coccoid forms. Cells 2.0 microns long by 0.5 to 0.7 micron broad; 3.0 to 5.0 microns long in division stages. Coccoid forms stained, 0.5 to 0.7 micron long. Motile with a single polar flagellum. Gram stain not reported. Gelatin stab : Very slight surface growth after one month ; the streak then shows a beaded line. No liquefaction. Agar colonies : Surface colonies appear as a whitish growth in a depression, sur- rounded by a white ring. The colony is later surrounded by a ring of liquid agar. Deep colonies show a clear area and may be irregular, oval or angular. Agar slant : A deep groove is cut along the inoculation streak, whitish growth along sides. The gel is later much weak- ened. Broth: Slightly turbid. No pellicle. Acid from glucose, lactose and mal- tose. No acid from sucrose or glycerine. Utilizes ammonia salts as a source of nitrogen. Decomposes cellulose and agar. The presence of one per cent glucose pre- vents the liquefaction of agar. Nitrites produced from nitrates. Starch is hydrolyzed. Aerobic. Temperature relations: Optimum 25° C, will grow at 16° but not at 34 °C. Habitat: Soil. 18. Vibrio cyclosites Gray and Thorn- ton. (Gray and Thornton, Cent, fiir Bakt., II Abt., 73, 1928, 92.) From Greek kyklos, circle or ring; sited, to eat; M. L. cyclosites, feeding on rings, i.e. ring compounds. Curved rods: 0.5 to 1.0 by 1.5 to 4.0 microns. Motile with a single polar flagellum. Gram-negative. Gelatin colonies : Circular, buff to brown, flat, smooth, glistening, entire. Gelatin stab: No liquefaction. Agar colonies : Circular to irregular, pale buff (later greenish), smooth, entire. Agar stab: Filiform, greenish buff, raised, smooth, undulate. FAMILY PSEUDOMOXADACEAE 201 Broth : Turbid. Indole not reported. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid from glucose. Attacks phenol and ?n-cresol. Aerobic, facultative. Optimum temperature 30° to 35 °C. Habitat : Soil. 19. Vibrio percolans Mudd and War- ren. (Jour, of Bact., 8, 1923, 447.) From Latin, percolo (percolatus) , filter- ing. Curved rods: 0.3 to 0.4 by 1.5 to 1.8 microns, occurring singly or in short chains. Pleomorphic. Actively motile by means of 1 to 3 polar flagella. Gram- negative. Gelatin stab: Xo liquefaction. Agar colonies : Circular, slightly con- vex, amorphous, entire. Agar slant : Bluish-white, glistening, streak. Broth: Turbid. Pellicle, sediment. Litmus milk : L'nchanged. Potato: White, slimy streak. Indole not formed. Nitrites not produced from nitrates. Blood serum not licjucfied. Starch not hydrolj'zed. No action on carbohydrates. Passes through bacterial filters. Aerobic, facultative. Optimum temperature 30°C. Non-pathogenic. Source: Isolated from haj^ infusion. Habitat: Water. 20. Vibrio andoi Aoi and Orikura. (Cent. f. Bakt., II Abt., 74, 1928, 331.) Named for Andoi, a Japanese scientist. Curved rods, with more or less tapering ends, c- or s-shaped, 0.5 to 0.8 by 1.5 to 2.5 microns. Motile, with a single polar flagellum. Gram-negative. Gelatin: No growth. Agar media : No growth. Broth: No growth. Litmus milk : No growth. Potato : No growth. Ammonium sulfate agar colonies : Punctiform, circular, concave, surrounded with clear zone. Ammonium sulfate agar slant : Grayish, becoming straw-yellow, sinking into the medium as the agar liquefies. Cellulose media: No growth. Starch hydrolyzed. Glucose, fructose, galactose, mannose, xylose and "honyak" are fermented. Xylan is decomposed. Cellobiose is decomposed. Aerobic, facultative. Optimum temperature 25 °C. Mini- mum 8 °C . Maximum 37 °C . Source : Rotted stable manure. Habitat: Presumably decomposing or- ganic matter. 21. Vibrio fetus Smith and Taylor. (Spirillum causing abortion in sheep, ^lacFadyean and Stockman, Rept. Dept. Comm. Ministr\' Agric. on Epizootic Abortion, London, 1909, 156; ibid., 1913, 111 ; Spirillum associated with infectious abortion, Smith, Jour. Exp. Med., 28, 1918, 701; Smith and Taylor, ihid., 30, 1919, 299; Spirillum fetus Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 552; Vibrio foetus ovis Buxton, First Rept. of Director, L^niv. Cambridge Instit. Animal Pathol., 1929-1930, 47.) From li. foetus, fetus. Curved rods : The smallest forms ap- pear as minute curved s-shaped lines, other forms very long; 0.2 to 0.5 by 1.5 to 5.0 microns. Motile by means of one, rarely two, polar flagella. Occasionally forms capsules. Granules present in older cultures. Gram-negative. Gelatin: Xo growth. Agar slant: No surface growth by freshly isolated strains. Laboratory strains produce a scanty, grayish-white, glistening surface growth. Subsurface agar colonies : Small, yel- low, opaque. Broth : A viscid ring pellicle may ap- pear, faint clouding of the medium occurs ; a filmy, stringy deposit may settle out. Litmus milk: No growth 202 MANUAL OF DETERMINATIVE BACTERIOLOGY Potato: No growth. Indole not formed. Nitrite production not reported. Blood serum slant: Feeble growth. No liquefaction. No gas from carbohydrates. No change or slightly acid from glucose, lactose and sucrose. Optimum temperature 37°C. With- stands 55°C for 5 minutes. Aerobic or microaerophilic. Pathogenesis : Causes abortion in cattle. Source : Twenty-two strains isolated from the placentas or fetuses of cows having abortion. Habitat: Causes abortion in cattle. 22. Vibrio pierantonii (Zirpolo) Meiss- ner. {Bacillus pierantonii Zirpolo, Boll. Soc. nat. Napoli, 30, 1918, 206; Meissner, Cent. f. Bakt., II Abt., 67, 1926, 200.) Named for Pierantoni, an Italian bac- teriologist. Rods : 0.5 by 1.5 microns, with rounded ends. Motile with one to three polar flagella. Gram-negative. Gelatin colonies: Circular, and irregu- larly lobulate. Gelatin stab: No liquefaction. Agar colonies: Circular, light green, smooth, entire. Glycerin agar slant : Slightl}^ luminous streak. Broth : Turbid, with pellicle. Indole not formed. Acid from glucose and maltose. Some strains also attack lactose, sucrose and mannitol. Best growth in alkaline media. Aerobic, facultative. Optimum temperature 37 °C. Source : Isolated from the photogenic organ of the cephalopod Sepiola inter- media Naef . Appendix:* The following species have also been listed in the literature. Many are inadequately described. Microspira bonhojjli Migula. (Bonhoff , Arch. f. Hyg., 19, 1893, 252; Migula, Syst. d. Bakt., 2, 1900, 1008.) From water. Microspira canalis Migula. {Spiril- lum saprophiles y and Vibrio saprophiles 7 Weibel, Cent. f. Bakt., 2, 1887, 469; Migula, Syst. d. Bakt., 2, 1900, 1004; Microspira cloaca Chester, Man. Determ. Bact., 1901, 341.) Possibly identical with Microspira saprophiles Migula, Microspira weibelii Migula, Vibrio siirati Ford, Vibrio smithii Ford. From sewage. Microspira coprophila Migula. (Group 3, No. 6, Kutscher, Ztschr. f. Hyg., 19, 1895, 475; Migula, Syst. der Bakt., 2, 1900, 986.) From fecal matter. Microspira maasei (v. Hoff) Migula. {Spirillum maasei v. Hoff, Cent. f. Bakt., I Abt., 21, 1897, 797; Migula, Syst. d. Bakt., 2, 1900, 978.) Possibly a variety of Vibrio comma Winslow et al. From Rotterdam tap water. Microspira milleri Migula. (Miller, Deutsche med. Wchnschr., 11, 1885, 138; Migula, Syst. d. Bakt., 2, 1900, 981; Spirillum milleri Holland, Jour. Bact., 5, 1920, 225; Vibrio milleri Holland, ibid.) Probably identical with Vibrio proteus according to Migula. From dental caries. Microspira rmirmanensis Issatchenko. (Recherches sur les microbes de I'ocean glacial arctique (in Russian), Petrograd, 1914, 240.) From sea water. Microspira saprophiles Migula. (Heu- vibrio 13, Weibel, Cent. f. Bakt., 2, 1887, 469 ; Vibrio saprophiles /3 Weibel, Cent, f . Bakt., 4, 1888, 225; Migula, Syst. d. Bakt., 2, 1900, 1006; Microspira weibell Chester, Man. Determ. Bact., 1901, 230.) Probably identical with Microspira cloaca Chester and Vibrio surati Ford. From sewage. Microspira tyrosinalica Beijerinck. (Kon. Akad. Wetenschappen, Amster- dam, 13, 1911, 1068.) From sewage. Microspira weibelii Migula. {Vibrio * Prepared by Mr. Wm. C. Ilaynes, New York State Experiment Station, Geneva, New York, Jan., 1939; Revised by Capt. Wm. C. Haynes, Sn. C, Fort Bliss, Texas, July, 1943. FAMILY PSEUDOMONADACEAE 203 saprophiles a Weibel, Cent. f. Bakt., 2, 1887, 465; ibid., 4, 1888, 225; Migula, Syst. d. Bakt., 2, 1900, 1005; Microspira saprophilc Chester, Manual Determ. Bact., 1901,341; V ibrio saprophiles Ford, Textb. of Bakt., 1927, 356.) Possibly identical with Microspira cloaca Chester, Vibrio surati Ford. ]' ibrio smithii Ford. From sewage. Spirillum lipofcrutn Beijerinck. (Cent. f. Bakt., II Abt., 63, 1925, 353; Chromatium lipoferum Bergey et al., Manual, 3rd ed., 1930, 531.) From garden (>arth and sewage. Giesberger (Beitriige zur Kenntnis der Gattung Spirillum Ehbg., Inaug. Diss., Delft. 1936, 64) regards this organism as a Vibrio. Has a single polar flagellum. SpiriUurn nasicola Trevisan. (Xasen- schleimvibrio, Weibel, Cent. f. Bakt., 2, 1887, 465; Trevisan, I generi e le specie delle Batteriacee, 1889, 24; Vibrio nasalis Eisenberg, Bakt. Diag.. 3 Aufl., 1891, 213; Spirillum nasale Sternberg, Man. of Bact., 1893, 697; Spirosoma nasale Migula, in Engler and Prantl, Die nattirl. Pflanzenfam., 1. I a, 1895. 31.) From human nasal mucus. Spirillum parvum Esmarch. (Cent. f. Bakt., I Abt., Orig., 32, 1902, 565; also see Zettnow, ibid., 78. 1910, 1; Vibrio j'>arvus Lehmann and Neumann, Bakt. Diag., 4 Aufi., 2, 1907. 494.) From de- caying organic matter. Vibrio albensis Lehmann and Neu- mann. (Elbe vibrio, Dunbar, Deutsch. raed. Wochnschr., 19, 1893, 799; Leh- mann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 340; Microspira dunbari Migula, Syst. d. Bakt., 2, 1900, 1013; Photospiril- lum dunbari Miquel and Cambier, Traite de Bact., Paris, 1902, 881 ; Photobacterium dunbari Ford, Textb. of Bakt., 1927, 621 .) From water of the river Elbe. Phos- phorescent. Vibrio amylocella Gray. (Canad. Jour. Res., 17, 1939, 154.) Decomposes cel- lulose. Produces glucose from starch. From soil. Vibrio anguillarum Bergman. (Ber. a. d. k. Bayr. Biolog. Versuchstat., Mun- chen, 2, 1909.) From an infectious dis- ease of eels. Vibrio aureus Weibel. (Weibel, Cent, f. Bakt., 4, 18S8, 225, 257, 281; Spirillum aureum Trevisan, I generi e le specie delle Batteriacee, 1889, 24; Spirillum aureum Sternberg, Man. of Bact., 1893, 700; Spirosoma aureum Migula, Syst. d. Bakt.. 2, 1900, 958.) Possibly identical with Vibrio flavns Weibel and Vibrio flavescens Weibel. From sewage. Vibrio beijerinckii Stanier. (Jour. Bact., 42, 1941, 527-554.) Marine agar- digesting vibrio. Vibrio buccalis Prevot. (Vibrion B, Repaci, Compt. rend. Soc. Biol., Paris, 1909, 630; Prevot, :Man. de Classif. des Bact. Anacr., Paris, 1940. 82. i Anaerobe. From the buccal cavitJ^ Vibrio bulbosa Kalnins. (Latvijas Universitates Raksti, Serija I, No. 11, 1930,237.) Decomposes cellulose. From soil. Vibrio cardii Klein. (Cent. f. Bakt., I Abt., Orig., 38, 1905, 173.) Possibly identical with Vibrio cuneatus Gray and Thornton and ]' ibrio marinus Ford. From the mussel (Cardium edule). Vibrio castra Kalnins. (Latvijas Uni- versitates Raksti, Serija I, No. 11, 1930, 241.) Decomposes cellulose. From soil. Vibrio choleroides a and /3 Bujwid. (Cent. f. Bakt., 13, 1893, 120; Microspira choleroides ]\Iigula, Syst. d. Bakt., 2, 1900, 992.) Probably a less vigorous strain of Vibrio comma Winslow et al. according to Chester, Man. Determ. Bact., 1901, 337. From water. Vibrio chrxjsanthemoides Lehmann and Neumann. (Spirillum-like organism, Jones. Cent. f. Bakt., II Abt.. 14. 1904, 459; Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907. 493.) From five samples of tap water and sewage. Vibrio crassus (Veillon and Repaci) Prevot. {Spirillum crassum Veillon and Repaci, Ann. Inst. Past., 26, 1912, 306; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 85.) Anaerobe. From the buccal cavity. Vibrio crassus var. D, Prevot. (Spirille 204 MANUAL OF DETERMINATIVE BACTERIOLOGY D, Repaci, Ann. Inst. Past., 26, 1912, 550; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 86.) Anaerobe. From the buccal cavity. Vibrio crasteri Hauduroy et al. (Craster, in Violle, Le Cholera, Masson edit., 1919; Hauduroy et al.. Diet. d. Bact. Path., 1937, 541.) Isolated from healthy persons. Resembles Vibrio comma. Vibrio cucumis Kalnins. (Latvijas tJniversi tales Raksti, Serija I, No. 11, 1930,243.) Decomposes cellulose. From soil. Vibrio devorans Beijerinck. (Cent. f. Bakt., II Abt., 11, 1903, 598.) From water. Vibrio drennani Chalmers and Water- field. (Drennan, Jour. Inf. Dis., 14, 1914, 251; Chalmers and Waterfield, Jour. Trop. Med., 19, 1916, 165.) Colon- ies white, turning dark brown. From feces. Vibrio flavescens Weibel. (Cent. f. Bakt., 4, 1888, 225, 257, 281; Spirillum flavescens Trevisan, I generi e le specie delle Batteriacee, 1889, 24; Spirillum, flavescens Sternberg, Man. of Bact., 1893, 700; Spirosoma flavescens Migula, Syst. d. Bakt., 2, 1900, 959.) Possibly identical with Vibrio aureus Weibel and Vibrio flavus Weibel. From sewage. Vibrio flavus Weibel. (Cent. f. Bakt., 4, 1888, 225, 257, 281; Spirillum flavum Trevisan, I generi e le specie delle Bat- teriacee, 1889, 24; Spirillum flavum Sternberg, Man. of Bact., 1893, 700; Spirosoma flavumM\g\i\&, Syst. d. Bakt., 2, 1900, 959.) Possibly identical with Vibrio aureus Weibel and Vibrio flaves- cens Weibel. From sewage. Vibrio fuscus Stanier. (Jour. Bact., 42, 1941, 540.) Marine agar-digesting vibrio. Vibrio gauducheau Hauduroy et al. (Gauducheau, in Violle, Le Cholera, Masson 6dit., 1919; Hauduroy et al., Diet. d. Bact., 1937, 543.) From the blood of a fever patient. Resembles Vibrio comma. Vibrio ghinda Pfeiffcr. (Pasquale, Gior. med. d. r. esercito, 1891 ; Pfeiffer, in Fliigge, Die Mikroorganismen, 2, 1896, 590; Microspira ghinda Migula, Syst. d. Bakt., 2, 1900, 996.) From water. Vibrio grossus (Migula) Ford. (Vibrio No. 1, Kutscher, Ztschr. f. Hyg., 20, 1895, 46 ; Microspira grossa Migula, Syst. d. Bakt., 2, 1900, 1012; Ford, Te.xtb. of Bact., 1927, 343.) From liquid manure. Vibrio halobicxis desulfuricans Horo- witz-Wlassowa and Sonntag. (Arb. a. d. Staatl. wissensch. Nahrungsmittel- Institut 1931 (Russian); see Ztschr. f. Unters. d. Lebensm., 62, 1931, 597.) A halophilic vibrio found in salted sar- dines, anchovies and other marine fish. Vibrio helcogenes Fischer. (Cent. f. Bakt., 14, 1894, 73; Microspira helcogenes Migula, Syst. d. Bakt., 2, 1900, 978.) From descriptions, indistinguishable from Vibrio proteus according to Chester, Man. Determ. Bact., 1901, 339. From feces. Vibrio hyos Ford. (Vibrio No. 3, Kutscher, Ztschr. f. Hyg., 20, 1895, 46; Spirillum mobile Migula, Syst. d. Bakt., 2, 1900, 1020; Ford, Textb. of Bact., 1927, 342.) Isolated from liquid manure. Vibrio iners Besson, Ranque and Senez. (Compt. rend. Soc. Biol. Paris, 79, 1918, 1097.) From the feces of persons having dysentery. Vibrio intermedins (Migula) Ford. (Group V, No. 9 of cholera-like vibrios, Kutscher, Ztschr. f. Hyg., 20, 1895, 481; Microspira intermedia Migula, Syst. d. Bakt., 2, 1900, 967; Ford, Textb. of Bact., 1927,342.) Possibly identical with Vib- rio berolinensis Neisser. From water. Vibrio ivanoff Pfeiffer. (Ivanoff, Ztschr. f. Hyg., 15, 1893, 134; Pfeiffer, in Fliigge, Die Mikroorganismen, 2, 1896, 592.) Probably a variety of Vibrio comma Winslow et al. according to Chester, Man. Determ. Bact., 1901, 337. From feces of a cholera patient. Vibrio jejuni Jones, Orcutt and Little. (Jour. Exp. Med., 53, 1931, 853.) From small intestine of calves suffering from diarrhea. Vibrio l-egallcnsis Hauduroy et al. FAMILY PSEUDOMONADACEAE 205 (Diet. d. Bact. Path., 1937, 544.) From water. Vibrio klimenko Hauduroy et al. (Klimenko, in Violle, Le Cholera, Masson edit., 1919; Hauduroy et al., Diet. d. Bact. Path., 1937, 544.) Resembles Vibrio comma. From the intestine. Vibrio leidensis Horst. (Inaug. Diss., Leiden. 1921; abst. in Cent. f. Bakt., I Abt., Ref., 73, 1922, 282.) From a liver abscess. *Vibrio lingualis P^isenberg. (Zungen- belag-Vibrio, Weibel, Cent. f. Bakt., 4, 1888, 227; ELsenberg. Bakt. Diag., 3 Aufl., 1891, 212; Spirillum linguae Sternberg, Alan, of Bact., 1893, 697; Spirosoma lin- guale Aligula, in Engler and Prantl, Die naturl. Pflanzenfam.. /, 1 a, 1895, 31.) From deposit on the tongue. Vibrio lissabonensis Pestana-Betten- court. (Cent. f. Bakt., 16, 1894, 401.) According to Chantemesse identical, or nearly so, with Vibrio proleus. From descriptions, indistinguishable from Vib- rio proteus according to Chester, Man. Determ. Bact., 1901, 339. From feces of a cholera patient. Vibrio malamoria Kalnin.s. (Latvijas U^niversitates Raksti, Serija I, No. 11, '930,250.) Decomposes cellulose. From soil. Vibrio marinus (Russell) Ford. {Spirillum marinum Russell, Ztschr. f. Hyg., 11, 1891, 165; Microspira marina Migula, Syst. d. Bakt., 2, 1900, 1002; Ford, Textb. of Bact., 1927, 347.) From sea water. Closely resembles Vibrio cuneatus Gray and Thornton and Vibrio cardii Klein. Vibrio tuassauah Pfeiffer. (Pasquale, Gior. med. d. r. esercito, 1891; Pasquale, Baumgarten's Jahresberichte,7, 1891,336; Pfeiffer, in Flligge, Die Mikroorganismen, 2, 1896, 589 ; Microspira massauah Migula, Syst. d. Bakt., 2, 1900, 963; Spirillum massauah Chester, Manual Determ. Bact., 1901, 343; Spirillum massowah Holland, Jour. Bact., 5, 1920, 225; Vibrio massowah Holland, ibid.) From feces of a cholera patient. Vibrio mulicris Prevot. (Man. de Classif. des Bact. Anaer., Paris, 1940, 84.) Anaerobe. From the female genital tract. Vibrio napi Kalninis. (Latvijas Uni- versitates Raksti, Serija I, No. 11, 1930, 252.) Decomposes cellulose. From soil. Vibrio n'dianka Hauduroy et al. (Thiroux, in Violle, Le Cholera, Masson edit., 1919; Hauduroy et al.. Diet. d. Bact. Path., 1937, 546.) Isolated from a patient having a cholera-like disease. Vibrio pericoma Kalnins. (Latvijas tJniversitates Raksti, Serija I, No. 11, 1930,256.) Decomposes cellulose. From soil. Vibrio polymorphus Prevot. (Spiro- chete B, Repaci, Ann. Inst. Past., 26, 1912, 544; Vibrio pseudospirochaeta B, Weinberg, Nativelle and Prevot, Les Microbes Anaerobies, 1936, 852; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 83.) Anaerobe. From the buccal cavity. Vibrio polymorphus var. peritriche Prevot. (Spirochete C, Repaci, Ann. Inst. Past., 26, 1912, 548; Vibrio pseudo- spirochaeta C, Weinberg, Nativelle and Prevot, Les Microbes Anaerobies, 1936, 854; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 84.) Anaerobe. From the buccal cavity. Vibrio portuensis (Migula) Ford. (Der portuenser Vibrio, Jorge, Cent. f. Bakt., I Abt., 19, 1896, 277; Microspira portuen- sis Migula, Syst. d. Bakt., 2, 1900, 1007; Ford, Textb. of Bact., 1927, 353.) From water. Vibrio prima Kalnins. (Latvijas Uni- versitates Raksti, Serija I, No. 11, 1930, 235.) Decomposes cellulose. From soil. Vibrio pseudospirochaeta Prevot. (Spirochete A, Repaci, Ann. Inst. Past., 26, 1912, 539 ; Vibrio pseudospirochaeta A, Weinberg, Nativelle and Prevot, Les Microbes Anaerobies, 1936, 849; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 83.) Anaerobe. From tlie buccal cavity. Vibrio putridus Prevot. (Vibrion C, Repaci, Compt. rend. Soc. Biol. Paris, See Nocardia lingualis Chalmers and Christopherson. 206 MANUAL OF DETERMINATIVE BACTERIOLOGY 1909, 630; Prevot, JMan. de Classif. des Bact. Anaer., Paris, 1940, 83.) Anaerobe. From the buccal cavity. Vibrio pyogenes (Doerr) Lehmann and Neumann. (Eiterspirillum, Mezinescu, Cent. f. Bakt., I Abt., Orig., 35, 1904, 201; Spirillum pyogenes Doerr, Cent. f. Bakt., I Abt., Orig., S8, 1905, 15; Leh- mann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 493.) From pus in a case of pyelitis calculosa. Non-motile. Vibrio ranicula Kalnins. (Latvijas tjniversitates Raksti, Serija I, No. 11, 1930,248.) Decomposes cellulose. From soil. Vibrio rigensis Kalnins. (Latvijas tjniversitates Raksti, Serija I, No. 11, 1930,254.) Decomposes cellulose. From soil. Vibrio rubicundus Gottron et al. (Gottron, Weaver and Sherago, Jour. Bact., 4S, 1942, 61.) From a trickling filter. Vibrio septicus Kolle. (Kolle and Schumann in Kolle and Wassermann, Handb. d. path. Mikroorg., 2 Aufl., 4, 1912, 101.) Identical with Vibrio comma culturally and morphologicalh*. From a cholera-like disease. Vibrio smithii (Migula) Ford. (Smith, Cent. f. Bakt., 10, 1891, 177; Microspira smithii Migula, Syst. d. Bakt., 2, 1900, 1006; Ford, Textb. of Bact., 1927, 340.) Possibly identical v,-ith Microspira sapro- philes Migula, Microspira weibelii Mi- gula, Microspira cloaca Chester and Vibrio surati Ford. From abscesses of large intestine of swine. Vibrio spermatozoides Loffier. (Cent, f. Bakt., 7, 1890. 638.) From kohlrabi infusions. Vibrio sputigenus (Miller) Prevot. {Spirillum sputigenum Miller, Die Mikro- org. d. Mundhohle, 2nd ed., 1892; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 85; not Vibrio sputigenus Bergey et al., Manual, 1st ed., 1923, 80.) An- aerobe. From the buccal cavity. Vibrio sputigenus var. minutissimus Prevot. (Muhlens, Cent. f. Bakt., I Abt., 48, 1909, 523; Pr6vot, Man. de Classif. des Bact. Anaer., Paris, 1940, 85.) Anaerobe. From the buccal cavity. Vibrio sputorum Prevot. (Man. de Classif. des Bact. Ana6r., Paris, 1940, 85.) Anaerobe. Isolated from a case of bronchitis. Vibrio stationis Kalnins. (Latvijas tjniversitates Raksti, Serija I, No. 11, 1930,239.) Decomposes cellulose. From soil. Vibrio stomatitis Prevot. (Vibrion A, Repaci, Compt. rend. Soc. Biol. Paris, 1909, 630; Prevot, Man. de Classif. des Bact. Anaer., Paris, 1940, 82.) Anaerobe. From the buccal cavity. Vibrio subtilissimus (Migula) Ford. (Spirillum No. 1, Kutscher, Ztschr. f. Hyg., 20, 1895, 46; Spirillum tenerrimum Lehmann and Neuman, Bakt. Diag., 2, 1896, ZA^; Spirillum subtilissimumMigula, Syst. d. Bakt., 2, 1900, 1020; Ford, Textb. of Bact., 1927, 341.) Regarded by Kutscher as being probably identical with the organism found by Smith (Cent, f. Bakt., 16, 1894, 324) in swine dung. Resembles Vibrio strictus. Vibrio suis Ford. (Vibrio No. 2, Kutscher, Ztschr. f. Hyg., 20, 1895, 46; Spirillum coprophilum Migula, Syst. d. Bakt., 2, 1900, 1019; not Microspira coprophila Migula, loc. cit., 986; Ford, Textb. of Bact., 1927, 341.) From liquid manure. Vibrio surati (Lamb and Baton) Ford. {Spirillum surati Lamb and Paton, Arch. Int. Med., 12, 1913, 259; Treponema surati Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 514; Ford, Textb. of Bact., 1927, 337.) Isolated from a case of vegetative endocarditis. Closely resembles Vibrio smithii Ford, Microspira weibelii Migula, Microspira saprophiles Migula and Microspira cloaca Chester. Vibrio synthetica Kalnins. (Latvijas tjniversitates Raksti, Serija I, No. 11, 1930,245.) Decomposes cellulose. From soil. Vibrio tenuis Veillon and Repaci. FAMILY PSEUDOMONADACEAE 207 (Ann. Inst. Past., 26, 1912, 300.) An- aerobe. From the buccal cavity. Vibrio terrigenus Giinther. (Cent. f. Bakt., 16, 1894, 746; Spirillum terri- genum Migula, Syst. d. Bakt., 2, 1900, 1017; Microspira terrigena Chester, Man. Determ. Bact., 1901, 341.) Closely re- lated to Vibrio tonsillaris Stephens and Smith. From soil. Vibrio tonsillaris Stephens and Smith. (Cent. f. Bakt., 19, 1896, 929; Microspira tonsillaris Migula, Syst. d. Bakt., 2, 1900, 1009.) Closely related to Vibrio terrigenus Giinther. From buccal cavity. Vibrio toulonensis Hauduroy et al. (Vibrion, Defressine and Cazeneuve, in Violle, Le Cholera, Masson edit., 1919; Hauduroy et al.. Diet. d. Bact. Path., 1937, 547.) From mussel beds in the bay of Toulon. Vibrio xylitica Kalnins. (Latvijas tJniversitates Raksti, Serija I, No. 11, 1930,232.) Decomposes cellulose. From soil. Genus II. Desulfovibrio Kluyver and van Niel.* (Cent. f. Bakt., II Abt., 04, 1936, 369; Sporovibrio Starkey, Arch. f. Mikrobiol., 9, 1938, 300.) From M. L. desulfo, an abbreviation of the poorly constructed word desulfofication, used to indicate reduction of sulfur compounds by bacteria; vibrio, vibrio. Slightly curved rods of variable length, usually occurring singly but sometimes in short chains which have the appearance of spirilla. Swollen pleomorphic forms are common. Actively motile by means of a single polar flagellum. Strict anaerobes which reduce sulfates to hydrogen sulfide. Found in sea water, marine mud, fresh water, and soil. The type species is Desulfovibrio desulfuricans (Beijerinck) Kluyver and van Niel. 1. Desulfovibrio desulfuricans (Beijerinck) Kluyver and van Niel. {Bacterium hydrosuljureum ponticum Zelinsky, Proc. Russ. Phys. and Chem. Soc, 25, 1893, 298; Spirillum desul- furicans Beijerinck, Cent. f. Bakt., II Abt., 1, 1895, 1: Bacillus desidfuricans Saltet, Cent. f. Bakt., II Abt., 6, 1900, 648; Microspira desulfuricans Migula, Syst. d. Bakt., 2, 1900, 1016; Kluyver and van Niel, Cent. f. Bakt., II Abt., 94, 1936, 369; Vibrio desulfuricans Holland, Jour. Bact., 5, 1920, 225; Sporo- vibrio desulfuricans Starkey, Koninkl. Nederland. Akad. v. Wetenschappen, Proc, 41, 1938, 425; also in Arch. f. Microbiol., 9, 19.38, 268.) From U. L. present part, desidfurico, sulfur re- ducing. Slightly curved rods, 0.5 to 1.0 by 1 to 5 microns, usually occurring singly but sometimes in pairs and short chains which cause them to look like spirilla. Swollen pleomorphic forms are common. Older cells appear black due to precipi- tated ferric sulfide. Actively motile, possessing a polar flagellum. Gram- negative. Stains readilj' with carbol fuchsin. Grows best in freshwater media. Fails to develop in sea water upon initial isolation. Produces opalescent turbiditj' in ab- sence of oxygen in mineral media enriched with sulfate and peptone. Media containing iron salts blackened. Bacteria found associated with pre- cipitated ferrous sulfide. Peptone-glucose agar colonies (in absence of air) : Small, circular, slightly raised, dull, entire, soft in consistency. Gelatin not liquefied. Peptone, asparagine, glycine, alanine, aspartic acid, ethanol, propanol, butanol, * Prepared by Dr. Claude E. .Jolla, California, Jan., 1943. ZoBell, Scripps Institution of Oceanography, La 208 MANUAL OF DETERMINATIVE BACTERIOLOGY glycerol, glucose, lactate, succinate and malate known to be utilized as hydrogen donors. Produces up to 500 ml. H2S per liter. Nitrites not produced from nitrates. Reduces sulfate to hydrogen sulfide. Also reduces sulfites, sulfur, thiosulfates and hyposulfites. Optimum pll 6 to 7.5, limits pH 5 to 9. Optimum temperature 25 to 30° C. Maximum 35 to 40° C. Anaerobic. Habitat: Soil, sewage, water. aestuarii (van nov. {Microspira 2. Desulfovibrio Delden) comb, aestuarii van Delden, Cent. f. Bakt., II Abt., 11, 1904, 81; Vibrio dcsulfur leans (halophilic strain) Baars, Over Sulfaat- reductie door Bakterien, Diss. Delft, 1930, 164 pp.) From Latin, aestuarium, estuary. Morphologically indistinguishable from Desulfovibrio desulfur leans described above, although it has a greater tendency to pleomorphism, and is slightly larger. Motile, possessing a polar flagellum. Gram-negative. Grows preferentially in media prepared with sea water or 3 per cent salt mineral solution enriched with sulfate and peptone. According to Baars {loc. clt.) the marine species can be acclimatized to tolerate hypotonic salt solutions but Rittenberg (Studies on Marine Sulphate- Reducing Bacteria, Thesis, Univ. of Calif., 1941, 115 pp.) was unable to con- firm this observation. Likewise Ritten- berg was unable to acclimatize D. aes- tuarii to tolerate temperatures exceeding 45° C or to produce endospores. Produces faint turbidity in absence of oxygen in sea water enriched with sulfate and peptone. Organisms most abundant in sediment. Agar colonies: Small, circular, slightly raised, darker centers, entire, soft consistency. Gelatin not liquefied. Peptone, asparagine, glycine, alanine, glucose, fructose, ethanol, butanol. glycerol, acetate, lactate and malate known to be utilized in presence of sulfate. Reduces sulfate to hydrogen sulfide. Also reduces sulfites, sulfur, thiosulfates and hyposulfites. Produces up to 950 ml. H2S per liter. Nitrites not produced from nitrates. Optimum temperature 25° to 30° C. Maximum 35° to 40° C. Optimum pH 6 to 8, limits pH 5.5 to 8.5. Anaerobic. Habitat : Sea water, marine mud, brine and oil wells. 3. Desulfovibrio rubentschickii (Baars) eomb. nov. {Vibrio rubent- schickii Baars, Over Sulfaatreductie door Bakterien, Diss. Delft, 1930, 164 pp.) Named for L. Rubentschick. Slightly curved rods, 0.5 to 1.0 by 1 to 5 microns, usually occurring singly, sometimes in pairs and short chains. Actively motile, possessing a polar flagellum. Gram-negative. Morpho- logically indistinguishable from Desul- fovibrio desulfuricans. Reduces sulfate to hydrogen sulfide. Also reduces sulfites, sulfur, thiosulfates and hyposulfites. Culturally and physiologically like D. desulfuricans except that D. rubent- schickii utilizes propionic acid, butyric acid, valeric acid, palmitic acid, stearic acid, galactose, sucrose, lactose and maltose. Anaerobic. Habitat : Soil and ditch water. Appendix : The following species has also been regarded as belonging in this genus. Vibrio thermodesulfurlcans Elion. (Cent. f. Bakt., II Abt., 63, 1924, 58); Vibrio desulfuricans (thermophilic strain) Baars, Over Sulfaatreductie door Bakterien, Diss. Delft, 1930, 164 pp.; Sporovibrio desulfuricans Starkey (Ko- ninkl. Nederland. Akad. u. Wetenschap- pen, Proc, 41, 1938, 425, also see Arch. f. FAMILY PSEUDOMONADACEAE 209 Microbiol., 5, 1938, 268.) A thermophilic produces endospores. However, spore- sulfate-reducing anaerobe which grows at formation appears to be the exception 30 to 65°C. and which, according to rather than the rule. The pleomorphic, Starkey, produces endospores. Elion peritrichous, sporogenous, sulfate- described Vibrio thcrmodesulfuricans reducer is more rod-like than the asporo- (Cent. f. Bakt., II Abt., 63, 1924, 58) genous cultures and many cells of the which grows at temperatures no lower sporogenous cultures are Gram-positive than 30 to 40°C. and has an optimum of ^^.^^^^^^^ asporogenous cultures of Desnl- 55°C. Morphologically it is much like r -i ■ i u- ■ r^ j.- ^ ,^ , , ,,. • , rx f ovibr 10 desulfuncans are {jvam-negative, Desulfoinbrio desulfuricans and D. aes- " n ,• , ■ > , ,. , ,, ^i .. , ,, , ., ,, , ■,- r • all oi which leaves a question whether the iuarii although the thermophilic form is ,r , • r. -77 , , J 1-1 1 i-i I sporogenous suliate-reducer is a Bacillus shorter, more rod-like, less motile and more pleomorphic. According to Baars «^' ^^ Desulfoinbrto. Rittenberg (htudies (loc. cit.), Vibrio thermodesulfuricam »" ^^^^"^e Sulfate-reducing Bacteria, Elion can be acclimatized to grow at Thesis, Univ. Calif., 1941, 115 pp.) was lower temperatures and it is found unable to adapt the marine sulfate re- abundantly in environments where the ducer to grow at low salinities or at high temperature has never been as high as temperatures, nor could it be induced 30 °C. This observation is confirmed by to form spores. Starkey (Arch. f. Microbiol., 9, 1938, Desulforibrio halohydrocarbonoclasti- 268) who found further that the thermo- cus Zobell (U. S. Patent Xo. 2.413,278; philic form found in nature or developed Science News Letter, Jan. 1 1, 1947.) b}' acclimatization to higher temperatures From oil bearing rocks. Genus III. Cellvibrio Winogradshy* (Ann. Inst. Pasteur, 4^, 1929. 577.) From M. L. cell, an abbreviation for cellulose; vibrio, vibrio. Long slender rods, slightly curved, with rounded ends, show deeply staining gran- ules which appear to be concerned in reproduction. Monotrichous. Most species produce a yellow or brown pigment with cellulose. Oxidize cellulose, forming oxy- cellulose. Growth on ordinary culture media is feeble. Found in soil. The type species is Cellvibrio ochraceus Winogradsky. Key to the species of genus Cellvibrio. I. Xo growth on glucose or starch agar. A. Ochre-yellow pigment produced on filter paper. 1. Cellvibrio ochraceus. II. Growth on glucose and starch agar. A. Poor growth on starch agar. 1. Cream-colored pigment which becomes brown with age is produced on filter paper. 2. Cellvibrio flavescens. B. Abundant growth on starch agar. 1. Scanty growth on glucose agar. a. Intense yellow pigment produced on filter paper. 3. Cellvibrio fulvus. 2. Abundant growth on glucose agar. a. X"o pigment produced on filter paper. 4. Cellvibrio vulgaris. * Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, Sept., 1937; no change. July, 1943. 210 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Cellvibrio ochraceus Winogradsky. (Ann. Inst. Pasteur, J!^3, 1929, 549, 601.) From Greek, ochra, yellow ochre; M. L. like ochre, yellow. Plump, curved rods with rounded ends, 2.0 to 4.0 microns long, rarely occurring as spirals. Chromatic granule frequently found in center. Motile with a single flagellum. Gram-negative. Produces diffuse, light ochre-colored, mucilaginous colonies on cellulose silica gel medium. No action or growth on plain agar. No growth on peptone, glucose, starch or tragacanth gum agar. Filter paper streaks : Entire paper colored ochre-yellow in 48 hrs. Aerobic, facultative. Optimum temperature 20 °C. Distinctive character : Rapid ochre- colored growth. Habitat : Soil. Disintegrates vegetable fibers. 2. Cellvibrio flavescens Winogradsky. (Ann. Inst. Pasteur, 43, 1929, 608.) From Latin, part. adj. oi flavesco, to turn yel- low or golden. Plump, curved rods, flexuous, with rounded ends, 0.5 by 2.5 to 5.0 microns. Shows metachromatic granules. Motile with a single flagellum. Gram-negative. Produces diffuse, cream-colored growth becoming brownish; mucilaginous colo- nies on cellulose silica gel medium. Good growth on peptone agar. Colo- nies 1 mm in 4 days. Grows poorly on glucose, starch and gum agars. Filter paper streaks : Almost as rapid in growth as Cellvibrio ochraceus and colors entire paper in 2 to 3 days. Aerobic, facultative. Optimum temperature 20 °C. Distinctive characters : Smaller, less curved rods that grow on a greater variety of media than Cellvibrio ochra- ceus, but do not attack cellulose as readily. Source : Isolated from a pile of old damp sawdust. Habitat : Soil. Disintegrates vegetable fibers. 3. Cellvibrio fulvus Stapp and Bortels. (Culture Y, Dubos, Jour. Bact., 15, 1928, 230; Stapp and Bortels, Cent. f. Bakt., II Abt., 90, 1934, 42.) From Latin, fulvus, reddish yellow. Slightly curved rods: 0.3 to 0.4 by 1.5 to 3.0 microns. Show involution forms. Motile by means of a single polar flagellum. Gram-negative. Cellulose is decomposed. Grows on filter paper with an intense egg-yellow color which in older cultures may deepen to rust brown. Glucose agar: Very scanty growth. Sucrose agar: Very slight growth. Maltose agar : Abundant yellow growth. Lactose agar: Fairly abundant yellow growth. Starch agar: Very abundant, bright yellow growth which later turns brown. Nutrient broth : No growth. Temperature relations: Optimum 25° to 30°C. Minimum 5°C. Maximum 32° to 35 °C. No growth at 37 °C. Thermal death point 39° to 40 °C. Aerobic. Source : Isolated from forest soil in Germany and from soil in the United States. Habitat : Widely distributed in soils. 4. Cellvibrio vulgaris Stapp and Bortels. (Culture Co, Dubos, Jour. Bact., 15, 1928, 230; Stapp and Bortels, Cent. f. Bakt., II Abt., 90, 1934, 44.) From Latin, vulgaris, common. Curved rods : 0.3 by 2.9 to 4.0 microns. Shows involution forms. Motile by means of a single polar flagellum. Gram- negative. Cellulose is decomposed. Grows on filter paper without the formation of pigment. Glucose agar: Abundant growth. No pigment. Sucrose agar : Abundant slightly yellow growth. FAMILY PSEUDOMONADACEAE 211 Maltose agar: Abunilant yellowish to 35°C. No growth at 37 °C. Thermal growth. death point 44° to 45°C. Lactose agar: Very heavy growth. Aerobic Starch agar : Very abundant yellowish „ t , , , , , bource : Isolated from forest soil in growth . Nutrient broth : No growth. Germany and from soils in the United Temperature relations: Optimum 25° states. to30°C. Minimum 5°C. Maximum 32° Habitat: Widely distributed in soils. Genus IV. Cellfalcicula W inogradshy .* (Ann. Inst. Pasteur, 43, 1929, 616.) From M. L. cell, an abbreviation for cellulose; Latin dim.., Jalcicula, a small sickle. Short rods or spindles, not exceeding 2.0 microns in length, with pointed ends, containing metachromatic granules. Old cultures show coccoid forms. Monotri- chous. Oxidize cellulose, forming o.xycellulose. Growth on ordinary culture media is feeble. Soil bacteria. The type species is Cellfalcicula viridis Winogradsky. 1. Cellfalcicula viridis Winogradsky. (Ann. Inst. Pasteur, iS. 1929. 616.) From Latin, viridis, green. Plump, small spindles, 0.7 by 2.0 microns, with rounded ends. Motile with a single flagellum. Gram-negative. Produces diffuse green, mucilaginous colonies on cellulose silica gel medium. Filter paper streaks : Rapid spreading growth colored green in 3 days at 30 °C. Hydrocellulose agar: Growth rapid, green; minute yellowish-green, mucous colonies on streaking. No growth on peptone, glucose, starch or gum agar. Aerobic, facultative. Optimum temperature 20°C. Habitat : Soil. 2. Cellfalcicula mucosa Winogradsky. (Ann. Inst. Pasteur, 43, 1929, 621.) From Latin, viucosus, mucous. Plump, curved spindles, with slightly pointed ends. Motile with a single polar flagellum. Contain a single chromatic granule. Gram-negative. Produces diffuse, cream-colored, muci- laginous colonies on cellulose silica gel medium. Hydrocellulose agar : Abundant grayish growth. No growth on peptone, glucose, starch or gum agar. Aerobic, facultative. Optimum temperature 20°C. Habitat: Soil. 3. Cellfalcicula fusca Winogradsky. (Ann. Inst. Pasteur, 43, 1929, 622.) From Latin, fuscus, dark, tawny. Plump, curved spindles, 0.5 by 1.2 to 2.5 microns, with slightly pointed ends and a central chromatic granule. Motile with a single polar flagellum. Gram- negative. Produces diffuse, brownish, slightly marbled or veined colonies on cellulose silica gel medium. Filter paper streak : Paper becomes a partially transparent, dry, non-muci- laginous pellicle adhei-ent to gel. Aerobic, facultative. Optimum temperature 20 °C. Source : Isolated from a pile of old damp sawdust. Habitat: Probably rotting wood. * Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva i^ew \ork, Sept., 1937; no change, July, 1943. 212 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus V. Thiospira Vislouch-* (Jour, de Microbiologie, 1, 1914, 50; Sidjospir ilium Kluyver and van Niel, Cent. f. Bakt., II Abt., 9J^, 1936, 396.) From Greek, theion, sulfur; speira, coil. Colorless, motile, slightly bent rods, somewhat pointed at the ends, with granules of sulfur within the cells and a small number of flagella at the ends. The type species is Thinspira winograchkyi (Omelianski) Vislouch. 1. Thiospira winogradskyi (Ome- 2. Thiospira bipunctata (Molisch) lianski) Vislouch. {ThiospiriUum wi7io- Vislouch. (Spirillum bipunctatum Mo- gradskyi Omelianski, Cent. f. Bakt., II Hsch, Cent. f. Bakt., II Abt., 33, 1912, Abt., 14, 1905, 7m ; ThiospiriUum. granu- 55; Vislouch, Jour, de Microbiologie latum Molisch, Cent. f. Bakt., II Abt., (Russian), 1, 1914, 50.) From Latin, bi, 33, 1912, 55; Vislouch, Jour, de Micro- two; punctum, points, biologic '(Russian), 1, 1914, 50; Sulfo- Small, slightly bent sulfur spirilla, spirillum winogradskyi Kluyver and van markedly pointed at the ends, 6.6 by 14 Niel, Cent. f. Bakt., II Abt., 94, microns long 1.7 to 2 4 microns wide (in ,„„„ „„_ , T-T J i- ^^T■ J 1 the center of the cell). Both ends are 1936, 397.) Named for Wmogradsky, -xi i i i- ^ . , . , . , filled more or less with large volutin the Russian bacteriologist. (metachromatic) granules. Several mi- Large, sulfur spirilla, somewhat ^^^^^ granules of sulfur are present in the pointed at the ends, 2 to 2.5 microns ^j^^^. ^^^^^^ ^^^ sometimes at the ends, thick, to 50 microns long. Numerous qjj ^gjjg possess one flagellum at each granules of sulfur. Very motile, with end; young cells have a flagellum at one to two polar flagella. one end. Habitat: Curative mud. Habitat: Sea and salt waters. Gcnvs VJ. Spirillum Ehrenberg .^ (Ehrenberg, Abhaudlungon d. Berl. Akad., 1830, 38; Spirosoma Migula, Arb. bakt. Inst. Karlsruhe, 1, 1894, 237; Dicrospirillum Enderlein, Sitzber. Gesell. naturf. Freunde, Berlin, 1917, 313.) From Greek, speira, a spire or coil. Cells form either long screws or portions of a turn. Volutin granules are usually present. Usually motile by means of a tuft of polar flagella (5-20) which may occur at one or both ends of the cells. Aerobic, growing well on ordinary culturemedia, except for one saprophyte and the pathogenic species. These have not yet been cultivated. Usually found in fresh and salt water containing organic matter. The type species is Spirillum xindula (Miiller) Ehrenberg. Key to the species of genus Spirillum. I. One micron or less in diameter. 1. Volutin granules present. a. Slow to rapid liquefaction of gelatin, b. Grayish to brown growth on potato. 1. Spirillum vndula. * Prepared by Prof. D. H. Bergey, Philadelphia, Penn., October, 1922. t Revised by Prof. D. H. Bergey, Philadelphia, Pennsylvania, April, 1937; further revision by Prof. Robert S. Breed, New York State E.xperiment Station, Geneva, New York, June, 1943, based on Monograph by Giesberger, Inaug. Diss., Utrecht, Nov. 30, 1936. FAMILY PSEUDOMOiVADACEAE 213 bb. Light yellowish-orange growth on potato. 2. Spirillum serpens. aa. No liquefaction of gelatin. Of small size (0.5 micron in diameter). b. Colonies on agar white becoming brownish black and slightly wrinkled. 3. Spirillum itersonii. bb. Colonies on agar white and smooth. 4. Spirillum tenue. 2. No volutin granules observed. b. Single flagellum. 5. Spirillum virginianum. bb. Tuft of flagella. 6. Spirillum minus. II. Over one micron in diameter. 1. Grows poorly on peptone agar and potato. 7. Spirillum kutscheri. 2. Xot positively known to have been cultivated on artificial media. Very evident volutin granules. S. Spirillum volulans. 3. Cells more or less deformed by fat drops. 9. Spirillum lipoferum. 1. Spirillum undula rMiiller) Ehren- berg. {Vibrio undula MtiUer, Animal- cula infusoria et marina, 1786 ; Ehrenberg, Infusionstierchen, 1838; Spirillum undula minor Kutscher, Cent. f. Bakt., I Abt., 18, 1895, 614.) From Latin, undulatus, wave-like. Stout threads, 0.9 micron in diameter, with one-half to three turns. The wave lengths are 6 microns. Width of spiral, 3.0 microns. Tufts of three to nine flagella at each pole. Volutin granules present. Gram-negative. Gelatin colonies : The surface colonies are circular, granular, greenish-yellow, entire. Gelatin stab: Thick, white, rugose surface growth. Very slow liquefaction. Agar colonies: Grayish-white, smooth. Broth : Turbid. Potato: Grayish-brown growth. Indole not formed. Catalase positive. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25 °C. Cohn (Beitrage z. Biol. d. Pflanzen, /, Heft 2. 1875, 132) reports that he could not distinguish this organism from Vibrio prolifer Ehrenberg. Habitat: Putrid and stagnant water. 2. Spirillum serpens (MuUer) Winter. {Vibrio serpens Miiller, Animalcula in- fusoria et marina, 1786, 43; Winter, in Rabenhorst's Kryptogamen-Flora, 1, Die Pilze, 1884, 63.) From Latin, serpens, serpent. Long, curved rods with two to three wave-like undulations, 0.8 to 1.0 micron in diameter; wave length, 8 to 9 microns. Width of spiral 1.5 to 1.8 microns. Vo- lutin granules in cytoplasm. Motile, possessing tufts of flagella at both poles. Gram-negative. Gelatin colonies : Yellowish to brown- ish, granular, entire. Gelatin stab : Yellowish surface growth. Slow liquefaction. Agar colonies : Heavj- cream-colored growth. Agar slant: Grayish, with yellowish center, granular, entire. Broth: Turbid. Litmus milk : Unchanged. Potato: Clear orange-yellow growth Indole not formed. 214 MANUAL OF DETERMINATIVE BACTERIOLOGY Catalase positive. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 35 °C. Habitat : Stagnant water. 3. Spirillum itersonii Giesberger. (Inaug. Diss., Utrecht, 1936, 46 and 57.) Named for van Iterson, the Dutch bac- teriologist. The smallest of the spirilla isolated from water. First observed by van Iterson (Proc. Kon. Akad. v. Wetensch. Amsterdam, 5, 1902, 685). Small spirals, 0.5 micron in diameter. Wave length, 3 to 3.5 microns. Spiral width, 1 to 1.5 microns. Motile with bipolar tufts of flagella. Gram-negative. Grows readily on peptone agar. White colonies becoming brownish black, and slightly wrinkled. Gelatin stab : No liquefaction. Brownish-orange growth on potato. Volutin granules may be present. Catalase is i)roduced. Acid from glucose, fructose, ethyl alcohol, n-propyl alcohol, n-butyl alcohol, and glycerol. Utilizes acetic, propionic, n-butyric, tartaric, fumaric, lactic, citric, and succinic acids. Grows well in peptone broth. Also utilizes ammonia compounds. Anaerobic growth in the presence of nitrates when organic or ammonia nitro- gen is also available. Optimum temperature : 30°C. Source : Isolated from water. Habitat: Water. 4. Spirillxun tenue Ehrenberg. (In- fusionstierchen, 1838; see Bonhoff, Arch, f. Hyg., ^6, 1896, 162.) From Latin, tenuis, thin. Slender spirals. Diameter 0.7 micron. Wave lengths 4.5 to 5.0 microns. Width of spiral 1.5 to 1.8 microns. Actively motile in peptone water with tufts of flagella at each pole. Volutin granules present. Gram-negative. Agar colonies: White, smooth. Peptone agar slant : Heavy growth. Gelatin stab : No liquefaction. Catalase positive. Potato : Light brown growth. Acid from glucose and fructose. Slight acid from several other sugars and glycerols. Utilizes salts of acetic, pro- pionic, n-butyric, tartaric, lactic, citric, malic, and succinic acids. Ammonia compounds are used as a source of nitrogen. Optimum temperature, 30 °C. Source : Found in putrefying vegetable matter. Habitat: Putrefying materials. 5. Spirillum virginianum Dimitroff. (Jour, of Baet., 1:2, 1926, 19.) From M. L. genitive of Virginia. Spirals consisting of J to 3 complete turns in young cultures, older cultures showing 7 turns. 0.6 to 0.9 by 3 to 11 microns. Motile with a single polar flagellum on one or both ends. Gram- negative. Gelatin colonies: Entire, convex, cir- cular, moist, colorless. Gelatin stab : Growth along entire stab. No liquefaction. (Dimitroft", loc. cit.) Active liquefaction. (Giesberger, Inaug. Diss., Utrecht, 1936, 65.) Agar colonies: Dew drop, convex, entire, moist, colorless. Agar slant : Dew drop, isolated colonies. Broth: Cloudy, no flocculation. Uschinsky's protein-free medium : Abundant growth. Litmus milk: No growth. Loeffler's blood serum : Convex, isolated dew drop colonies. No lique- faction. Lead acetate agar: No H2S. Voges-Proskauer and methyl red nega- tive. No volutin granules observed (Gies- berger, loc. cit., p. 60). Potato: No growth. Indole not formed. Nitrites not produced from nitrates. No acid or gas from carbohydrates.- FAMILY PSEUDOMONADACEAE 215 (Dimitroff, loc. cit.). Utilizes lactates and citrates (Giesberger, loc. cit.). Aerobic, facultative. Optimum temperature 35 °C. Source : Isolated from mud on an oyster shell. Habitat : Probably muddy bottom of brackish water. 6. Spirillum minus Carter. (Carter, Sci. Mem. Med. Officers Army India, 3, 1887, 45; Spirillum minor Carter, ibid.; Spirochaeta laverani Breinl and King- horn, Mem. Liverpool Sch. Trop. Med., 21, 1906, 55; Spirochaeta miiri.'i Wenyon, Jour. Hyg., 6, 1906, 580; Spirochaeta muris var. virginiana MacNeal, Proc. Soc. Exper. Biol, and Med., 4, 1907, 125; Spirochaeta muris var. galatziana Mezin- cescu, Compt. rend. Soc. Biol. Paris, 66, 1909, 58; Treponema muris Moore, Principles of Microbiology, 1912, 414; Spirochaeta morsus mnris Futaki, Takaki, Taniguchi and Osumi, Jour. Exp. Med., 25, 1917, 33; Spirochaeta petit Row, Ind. Jour. Med. Res., 5, 1917, 386; Spironema muris Noguchi, Jour. Exp. Med., 27, 1918, 584; Spirochaeta japonica, Dujarric de la Riviere, Ann. de Med., 5, 1918, 184; Spirochaeta morsusmuris Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 447; Spiroschaudinnia morsusmuris Castellani and Chalmers, ibid.; Spiro- chaeta sodoku Troisier, 1920, according to Pettit, Contribution a I'Etude des Spirochetides, Vanves, II, 1928, 231; Treponema japonicum Brumpt, Nouveau Traitc de Medecine, Paris, 4, 1922, 505; Treponema morsus muris Brumpt, ibid., 506; Treponema minor Brumpt, ibid., 507; Treponema laverani Brumpt, ibid., 507; Treponema sodoku Brumpt, ibid., 514; Spirochaeta pettiti Row, Jour. Trop. Med. and Hyg., 25, 1922, 364; Treponemella muris San Giorgi, Patho- logica rivista, 14, 1922, 461; Borrelia muris Bergey et al.. Manual, 2nd ed., 1925, 435; Spirillum minus var. morsus muris Ruys, Cent. f. Bakt., I Abt., Orig., 103, 1927, 270; Spirillum minus var. ?nuris Ruys, ibid.; Spironevia minor Ford, Textb. of Bact., 1927, 962; Spiro- nema laverani Ford, ibid., 963 ; Spironema muris var. virginiana Ford, ibid., 963; Spirella morsusmuris Noguchi, in Jordan and P'alk, Newer Knowledge Bact. and Immun., 1928, 497; Spirella muris Noguchi, ibid.) From Latin, minus, less. Description taken from Adachi, Jour. E.xp. Med., 33, 1921, 647 and Giesberger, Inaug. Diss., Delft, 1936, 67. Short thick cells : 0.5 by 3.0 microns, having 2 or 3 windings which are thick, regular and spiral. Actively motile by means of bipolar tufts of flagella. Gram- negative. Has not been cultivated on artificial media. Aerobic, facultative. Pathogenic for man, monkej^s, rats, mice and guinea pigs. This species is regarded by some as a spirochaete. Because of its habitat and wide distribution it has been described under many different names. It is possible that some of these names indi- cate varieties or even separate species. See Beeson (Jour. Amer. Med. Assoc, 123, 1943, 332) for important literature. Source : Found in the blood of rats and mice. Habitat : The cause of rat-bite fever. Widely distributed. 7. Spirillum kutscheri Migula. {Spi- rillum undula majus Kutscher, Cent. f. Bakt., I Abt., 18, 1895, 614; Migula, Syst. d. Bakt., 2, 1900, 1024.) Named for Kutscher, the German bacteriologist who first isolated the organism. Stout threads, 1.5 microns in diameter. Wave lengths 10.5 to 12.5 microns. Width of spiral, 3 to 4.5 microns. May lose their spiral form on continued cultivation. Motile with tufts of flagella at the poles. Gram-negative. Gelatin colonies: Transparent, round, surface colonies. Deep colonies, dark brown. 216 MANUAL OF DETERMINATIVE BACTERIOLOGY Gelatin stab : Slow liquefaction. Agar colonies grow poorly, granular. Deep colonies yellowish-green to dark brown . Agar slant : Delicate, transparent growth. Potato: Limited growth. Volutin present. Catalase positive. Utilizes malic and succinic acids. Grows well on peptone broth. Also utilizes ammonia compounds. Optimum temperature, 22° to 27 °C. Source : Isolated from putrid materials and liquid manure. Habitat : Putrefying liquids. 8. Spirillum volutans Ehrenberg. (Prototype, Vibrio spirillum Miiller, Animalcula infusoria, 1786; Ehrenberg, Die Infusionstierchen als Volkommene Organismen, 1838.) From M. L. volutin. Spirals 1 .5 microns in diameter. Wave length, 13 to 15 microns, width of spiral, 4 to 5 microns. The largest of the spirilla. Slightly attenuated ends. Motile, pos- sessing a tuft of ten to fifteen flagella at each pole. Dark granules of volutin in the cytoplasm. Gram-negative. Migula (Syst. d. Bakt., 2, 1900, 1025) reports that this species has not been cultivated on artificial media, and that the cultures so described by Kutscher (Ztschr. f. Hyg., 20, 1895, 58) are of a different species which Migula names Spirillum giganteum. Vahle (Cent. f. Bakt., II Abt., 25, 1910, 237) later describes the cultural characters of an organism which he regards as identical with Kutscher 's organism. Giesberger (Inaug. Diss., Delft, 1936, 65) saw what he felt was the true Spirillum volutans but could not cultivate it. Optimum temperature 35°C. Habitat: Stagnant water. 9. Spirillum lipoferum Beijerinck. (Azotobacter spirillum Beijerinck, Kon. Akad. Wetensch. Amsterdam, 30, 1923, 431 quoted from Giesberger, Inaug. Diss., Delft, 1936, 24; Spirillum lipo- ferum Beijerinck, Cent. f. Bakt., II Abt., 63, 1925, 353; Chromatium lipo- ferum Bergey et al., Manual, 3rd ed., 1930, 531.) From Greek, lipos, fat; Latin, fero, to bear. Curved cells with one-half to one spiral turn, containing minute fat droplets. These may deform the cells. Motile with lophotrichous flagella. Gram- negative. Calcium malate agar colonies: Circu- lar, small, transparent, dry. The malate is oxidized to calcium carbonate. Cells contain fat drops. Peptone agar colonies : More abundant development. Cells lack fat drops and are typically spirillum in form. Glucose peptone broth : Cells actively motile with large fat drops. Fixes atmospheric nitrogen in partially pure cultures, i.e., free from Azotobacter and Clostridium (Beijerinck, loc. cit.). Schroder (Cent. f. Bakt., II Abt., 85, 1932, 17) failed to find fixation of nitrogen when she used cultures derived from a single cell. Aerobic. Optimum temperature 22°C. Beijerinck regards this as a transitional form between Spirillum and Azotobacter. Giesberger {loc. cit., p. 64-65) thinks it a Vibrio . Habitat: Garden soil. Appendix:* The following additional species have been mentioned in the literature. Many are inadequately de- scribed. Some may not belong here. * Prepared by Mr. Wm. C. Haynes, New York State Experiment Station, Geneva, New York, Jan., 1939; Revised by Capt. Wm. C. Haynes, Sn. C, Fort Bliss, Texas, .July, 1943. FA^nLY PSEUDOMOXADACEAE 217 Spirella canis Duboscq and Lebailly. (Compt. rend. Acad. Sci. Paris, 15Jt, 1912, 835.) From the stomach of a dog. Spirillum amyliferum Van Tieghem. (Bull. Soc. botan. de France, 26, 1879, 65.) Said to produce spores. Ford (Textb. of Bact., 1927, 364) thinks this organism was probably a spirochaete because of its mode of division. Found in frog spawn fungus of sugar factories. Spirillum attenuatum Warming. (Om nogle ved Danmarks Kyster levende Bakterier. Kjobenhavn, 1876; Spiro- soma attenuatum Migula, Syst. d. Bakt., 2, 1900, 959.) Ford {loc. cit., 363) states that this incompletely described organ- ism would now be regarded either as a spirillum or as a spirochaete. From sea coast of Denmark. Spirillum cardiopyrogenes Sardjito. (Geneesk. Tijdschr. voor Xed. -Indie, 72, 1932, 1359; ibid., 73, 1933, 822.) From lilood of a patient with pericarditis. Spirillum colossus Errera. (Rec. trav. bot. Bruxelle, 5, 1902; Abst. in Cent. f. Bakt., II Abt., 9, 1902, 608.) A giant form isolated from brackish sea water. Probably the same as Spirillum volutans Ehrenberg. Spirillum concentricum Kitasato. (Cent. f. Bakt., 3, 1888, 73.) Found in putrefying blood. Spirillum crassum Veillon and Repaci. (Ann. Inst. Past., 26, 1912, 300.) De- scribed as having peritrichous flagella. From lung lesions in human tuberculosis. Spirillum endo par agog icum Sorokin. (Cent. f. Bakt., 1, 1887, 465.) Described as producing spores in old cultures. From rain water in bark of poplar tree. Spirillum giganteum ^ligula. {Spiril- lum volutans Kutscher, Ztschr. f. Hyg., 20, 1895, 58; Migula, Syst. d. Bakt., 2, 1900, 1025.) From putrefying liquids. Spirillum hachaizae Kowalski. (Cent, f . Bakt., 16, 1894, 324 ; Spirillum hachaizi- cum Kowalski, ibid., 324; Spirochaete hachaizae Castellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 316 ; Treponema hachaizae Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 495.) Found in feces of cholera patients and also of healthy individuals. Spirillimi kolkwitzii Vislouch. (Jour, de Microbiol. (Russian), 1, 1914, 50.) Spirillum leiicomelaenum Perty. (Zur Kenntniss kleinster Lebensformen. Berne, 1852. Also see Koch, Mitt. Kais. Gesundheitsamte, /, 1881, 48.) From stagnant water. Spirilhun monospora Dobell. (Quart. Jour. Micr. Sci., 52, 1908, 121.) De- scribed as producing spores. From large intestine of frogs and toads. Spirillum nigrum Rist. (These med., Paris, 1898; see Cent. f. Bakt., I Abt., 30, 1901, 299.) Strict anaerobe from pus. Spirillum ostreae Xoguchi. (Jour. Exp. Med., 3.^, 1921,295.) From oysters. Spirillum, periplaneticum Kunstler and Gineste. (Compt. rend. Soc. Biol. Paris, 61, 1906, 135.) From the intestine of the cockroach, Periplaneta amcricana. Spirillum pyogenes Mezincescu. (Cent. f. Bakt., I Abt., Orig., 35, 1904, 201 ; Spirochaeta pyogenes Blanchard, Semaine Med., 26, 1906, 1; Treponema pyogenes Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 511.) From a case of pyelitis calculosa. Spirillum, rappini De Toni and Trevi- san. (Spirochaete, Rappin, Contr. a I'Etude d. Bacter. de la Bouche a I'Etat normal, 1881, 68; De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1009.) From the stomach of a dog. Spirillum recti physeteris Beauregard. (Compt. rend. Acad. Sci. Paris, 125, 1897,255.) From ambergris. Spirillum rugula (Miiller) Winter. {Vibrio rugula Miiller, Animalcula infu- soria, 1786; Cohn, Beitrage z. Biol. d. Pflanz., 1, Heft 2, 1872, 178; Bonhoff, Arch. f. Hyg., 26, 1896, 162; Winter, Die Pilze, in Rabenhorst's Kryptogamen- Flora, 1884.) Prazmowski found spores, but it is not certain his cultures were pure. Bonhoff also observed spores, but 218 MANUAL OF DETERMINATIVE BACTERIOLOGY concluded that they were due to con- taminating organisms (Ford, Textb. of Bact., 1927, 360). From water. Spirillum sporiferum Migula. (Syst. d. Bakt., 2, 1900, 1028.) Produces spores. The spirals in which the spore formation is beginning are like Spirillum leucomelaenum Perty (Ford, loc. cit., 336). Giesberger (loc. cit., p. 60) places this and other so-called spore-forming spirilla in Sporospirillum Orla-Jensen (Cent. f. Bakt., II Abt., 23, 1909, 340). From a bean infusion. Spirillum sputigenum Miller. (Die Mikroorganismen der Mundhohle. Leip- zig, 1892; Deutsche med. Wchnschr., 33, 1906, 1 and 348. ) Hoffman and Prowazek (Cent. f. Bakt., I Abt., Orig., 4i, 1906, 741) claim that Spirillum sputigenum hasperitrichousflagella. Giesberger {loc. cit., 63) places this in Selenomonas Prowazek (Cent. f. Bakt., I Abt., Orig., 70, 1913, 36). Muhlens (Cent. f. Bakt., I Abt., Orig., 48, 1909, 525) reports 1 to 3 flagella, the majority of the organisms having apparently a single thick fiagellum (a bunch of flagella) on the concave side (Ford, Zoc. cz7., 367). Anaerobic. From the buccal cavity. Spirillum slomachi Lehmann and Neu- mann. {Spirillum Form a, 13, y, 5 Salomon, Cent. f. Bakt., I Abt., 19, 1896, 433; Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 362.) Found in stomach of dog, cat and rat. Paraspirillum vejdovskiiDoheW. (Arch, f. Protistenk., 24, 1911, 97.) Found only once in fresh water containing Oscilla- toria. Flagellate flexible spiral cells described as possessing a nucleus. This may be a protozoan. SpirohaciUus gigas Certes. (Bull. Soc. Zool. France, 14, 1889, 322; abst. in Ann. de Microgr., 2, 1889-1890, 137.) From water. Vibriothrix tonsillaris Tunnicliff and Jackson. (Organism from Actinomyces- like granules, Tunnicliff, Jour. Inf. Dis., 38, 1926, 366; Tunnicliff and Jackson, ibid., 46, 1930, 12.) From tonsillar granules. May be identical with Lepto- thrix asteroide Mendel and as a Gram- negative, anaerobe may belong in Bac- teroides according to Rosebuiy (Bact. Rev., 8, 1944, 202). Vibriothrix zeylanica (Castellani) Cas- tellani. {Spirillum zeylanicum Castel- lani Jour. Ceylon Branch Brit. Med. Assoc, 7, 1910, 5 and Philipp. Jour. Sci., 5, No. 2, Sect. B., Medical Sciences, July, 1910; Vibrio zeylanicus Castellani, 1913, Bacillus zeylanicus Castellani, 1913 and Vibriothrix zeylanica Castellani, 1917, quoted from Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 1069; Spirobacillus zeylanicus Castellani, Spagnuolo and Russo, Bull. Soc. Path. Exot., 11, 1918, 271.) Motile. Gram- negative. From cases of dysenteric enteritis in Ceylon. This is the type species of the genus Vibriothrix Castel- lani (see Castellani and Chalmers, loc. cit., 1068). FAMILY AZOTOBACTERIACEAE 219 FAMILY III. AZOTOBACTERIACEAE BERGEY, BREED AND MURRAY.* (Preprint, Manual, 5th ed., October, 1938, v and 71.) Cells without endospores. Relatively large rods or even cocci, sometimes almost yeast-like in appearance. The type of flagellation in this genus has been definitely established as peritrichous. Gram-negative. Obligate aerobes, usually growing in a film on the surface of the culture medium. Capable of fixing atmospheric nitrogen when provided with carbohydrate or other energ\^ source. Grow best on media deficient in nitrogen. Soil and water bacteria. There is a single genus. Genus I. Azotobacter Beijeririck. (Beijerinck, Cent. f. Bakt., II Abt., 7, 1901, 567; Azotomonas Orla-Jensen, Cent. f. Bakt., II Abt., 24, 1909, 444.) The definition is identical with that of the family. From Gr. azous, not living. French, azote, nitrogen; Gr. bakiron, rod, stick. The type species is Azotobacter chroococcu?7i Beijerinck. 1. Azotobacter chroococcum Beijer- inck. (Cent. f. Bakt., II Abt., 7, 1901, 567 and 9, 1902, 3; Bacillus azotobacter Lohnis and Hanzawa, Cent. f. Bakt., II Abt., 4^, 1914, 1; Bacillus chroococcus Buchanan, General Syst. Bact., Balti- more, 1925, 194.) From Gr. chroa, color; coccos, grain; M. L. sphere. According to Lohnis and Smith (Jour. Agr. Res., 23, 1923, 401) Azotobacter beijerinckii Lipman (New Jersey Agr. Exp. Sta. Rept., 25, 1904, 247), Azoto- bacter woodstownii Lipman (ibid.), Azoto- bacter smyrnii Lipman and Burgess (Cent, f. Bakt., II Abt., U, 1915, 504) and Azoto- bacter hilgardii Lipman (Science, 29, 1909, 941) are identical with Azotobacter chroococcum. Greene (Soil Sci., 39, 1935, 327) studied Azotobacter chroococ- cum and Azotobacter beijerinckii by chemical analyses and found the chemical composition of the cells to be practically identical, but different from that of Azotobacter vinelandii and Azotobacter agile. Smith (private communication) feels that Azotobacter beijerinckii is a non-pigmented rough strain of Azoto- bacter chroococcum. Grows in absence of organic nitrogen- Rods : 2.0 to 3.0 by 3.0 to 6.0 microns, occurring in pairs and packets and oc- casionally in chains. The cells show three or four refractile granules. The organisms are surrounded hj a slimy membrane of variable thickness, usually becoming brownish in older cultures, due possibly to the conversion of tyrosine to melanin. The coloring matter is insolu- ble in water, alcohol, ether and chloro- form. Motile bj' means of numerous peritrichous flagella (Hofer, Jour. Bact., 47, 1944, 415.) Gram-negative. Gelatin colonies: Very small, circular, yellow, granular, later becoming yel- lowish-brown. Gelatin stab: Only slight growth in the stab. No liquefaction. Mannitol agar stab : Gray, may become brownish. Nutrient broth: No growth even in the presence of glucose ; peptone utilized with difficulty. Litmus milk : Becoming clearer in 10 to 14 days. Potato: Glossy, barely visible, slimy * Revised by Dr. A. W. Hofer, New York State Experiment Station, Geneva, New York, June, 1938; further revision by Dr. A. W. Hofer, July, 1943. 220 MANUAL OF DETERMINATIVE BACTERIOLOGY to wrinkled; may become yellowish, brownish-yellow or chocolate brown. The organism fixes atmospheric nitro- gen and gives off CO2, utilizing glucose and sucrose. Other generally used car- bon compounds are fructose, maltose, mannitol, inulin, dextrin, galactose, arabinose, starch, glycerol, ethyl alcohol, acetate, butyrate, citrate, lactate, mal- ate, propionate and succinate. Nitrate : Improves growth in amounts less than 1 gm. per liter ; greater amounts are toxic. Fixes nitrogen moderately actively. Chemical analysis : Four-day cultures grown upon mannitol agar (Greene, 1935), when dried, are found to contain less than 0.5 per cent of hemicelluloses, less than 20 per cent of crude protein, less than 5 per cent of ash, and more than 30 per cent of lignin-like materials. The nitrogen fraction contains less than 1 per cent of amide nitrogen, less than 1 per cent of humin nitrogen and about 1 per cent of basic nitrogen. Aerobic. Optimum temperature 25°C. to 28°C. Distinctive characters : Inability to grow in peptone media, even in the presence of glucose; frequent occur- rence of a dark brown or black pigment. Source : Isolated from soil. Habitat: Occurs naturally in the majority of neutral or alkaline field soils. 2. Azotobacter agile Beijerinck. (Cent. f. Bakt., II Abt., 7, 1901,577.) From L. agilis, agile, quick. In studies on the chemical composition of cells Greene (Soil Sci., 39, 1935, 327) found Azotobacter vinelandii Lipman (New Jersey Agr. Exp. Sta. Rept., 24, 1903, 238) to be very similar to Azoto- bacter agile Beijerinck. Smith and Loh- nis (Jour. Agr. Res., 23, 1923, 401) agree and state furthermore that the two are identical; they believe also that Azoto- bacter vitreiim Lohnis and Westermann (Cent. f. Bakt., II Abt., 22, 1908, 234) is another synonym of Azotobacter agile. Smith (private communication) states that Azotobacter vitreum is a very weak growing, smooth strain of Azotobacter agile. Kluyver and van Reenen (Arch. Mikrobiol., 4, 1933, 299) feel that a dis- tinction should be made between Azoto- bacter agile and Azotobacter vinelandii. In regard to the former, Kluyver and van den Bout (Arch. Mikrobiol., 7, 1936, 263) suggest that it be further subdivided into Azotobacter agile and Azotobacter agile var. atypica, the latter referring to an Azotobacter agile form that fails to produce pigment. Rods: 4 to 6 microns in length, almost spherical. Actively motile by means of numerous peritrichous flagella (Hofer, loc. oil). Some strains are reported to be non-motile. Gram-negative. Grows in absence of organic nitrogen. Gelatin : No liquefaction. Mannitol agar colonies : Circular, gray- ish white, translucent with whitish center. Washed agar colonies : Show slight bluish-green fluorescence. Mannitol agar slant: Grayish, trans- lucent, fluorescent. Plain agar slant : Yellowish-white, smooth, glistening, translucent with opaque center. Broth: Turbid, with sediment. Litmus milk : Becoming clear in 10 to 14 days. Potato : Yellowish-white, slimy, be- coming yellowish-brown. In the presence of organic acids, a greenish or reddish pigment is formed. The organism fixes atmospheric nitro- gen actively, and gives off CO2. Aerobic. Chemical analysis : Four-day cultures grown upon mannitol agar (Greene, 1935), when dried, contain more than 4 per cent of hemicelluloses, more than 45 per cent of crude protein, more than 7 per cent of ash, and less than 4 per cent of lignin- like materials. The nitrogen fraction contains more than 1 per cent amide nitrogen, more than 1 per cent humin FAMILY AZOTOBACTERIACEAE 221 nitrogen, and 2 per cent or more of basic nitrogen. Optimum temperature 25 °C to 28 °C. Distinctive characters : Lack of a brown pigment ; occasional fluorescence ; growth in peptone broth containing glucose. Source : Originally isolated from canal water at Delft. Habitat: Occurs in water and soil. 3. Azotobacter indicum Starkey and De. (Soil Sci. 47, 337, 1939.) From L. indict/a, of India. Rods: Ellipsoidal, from U. 5 to 1.2 by 1.7 to 2.7 microns when grown on nitrogen free glucose agar. One of the distinctive characteristics is the presence of two large, round, highlj'' refractive bodies in the cells, one usually at each end. Mo- tile by means of numerous peritrichous fiagella (Hofer. loc. cit.). Gram-nega- tive. The organism grows slowly but in time produces large amounts of slime. Has high acid tolerance, since it grows from pH 3 to 9. Sucrose or glucose agar plates : Colonies are colorless, round, very much raised, and uniformly turbid, having much the appearance of heavy starch paste. Af- ter two weeks, a buff to light brown color develop;. Mannitol agar slant : Grows very poorly. Peptone agar slant with 0.5 per cent glucose: Limited grayish growth. Nutrient broth : No growth. Liquid media generally : Turbidity with some sediment. Fixes atmospheric nitrogen readily with either glucose or sucrose as source of energy. Aerobic. Optimum temperature: 30°C. Distinctive characters : Tolerance of acidity, wide limits of pH tolerated, abundant slime production, large glob- ules of fat within cells. Source : Soils of India. Habitat : Soils. Appendix I : The relationship of the following species to the species placed in .420- tobacler is not yet entirely clear. Genus Azotomonas Stapp. (Cent. f. Bakt., II Abt., 102, 1940, 18; not Azotomonas Orla-Jen.sen, Cent. f. Bakt., II Abt., 24, 1909, 484.) Rod to coccus-shaped aerobic bacteria, motile by means of 1 to 3 polar Hagella. Xo endospores. No fat-like reserve food granules in the cells. Form acid and gas from glucose, and other sugars and alcohols. Form indole. Chemo-heterotrophic. Many carbon compounds other than sugars used as sources of energy. Active in the fixation of atmospheric nitrogen. Live in soil. From Gr. azvus, not living. French, azote, nitrogen; Greek, monas, a unit; M. L. monad. The type species is Azotomonas insolita. Azotomonas insolita Stapp. (Ab- microns. Motile with one to three polar stracts of Communications, Third In- flagella. Gram-negative. ternat. Congr. for Microbiol.. Sect. VIII, 1939, 306; abst.in Proc. Soil Sci. Soc. of America, 4, 1939, 244; Cent. f. Bakt., II Abt., 102, 1940, 1.) From Latin insolilus, unusual . Gelatin: No liquefaction. Agar slant: Glistening white growth. Agar colonics: Flat, whitish, edge entire. Weakly fluorescent. Broth : Strong turbidity. Sediment. Coccoid rods: 0.6 to 1.2 bv 0.6 to 1.8 Pellicle. 222 MANUAL OF DETERMINATIVE BACTERIOLOGY Milk: No change. Potato : Growth somewhat dry, not slimy, dirty gray, spreading. Nitrites produced from nitrates. Fixes nitrogen. Ammonium salts utilized. Acid and gas from adonitol, arabinose, dextrin, glucose, galactose, glycerine, inositol, lactose, fructose, maltose, man- nitol, mannose, raffinose, rhamnose, sali- cin, sorbitol, starch, sucrose and xylose. Starch is hydrolyzed. Hydrogen sulfide produced. Optimum temperature 25° to 30 °C. Minimum 7° to 9.5°C. Maximum 48°C. Good growth at 37 °C. Thermal death point 60 °C. Limits of pH 3.3 to 9.5. Aerobic. Source : From a mixture of chopped cotton husks and rice hulls. Habitat: Soil. FAMILY- RmzaMACH^E-- 223 FAMILY IV. RHIZOBL\CEAE COXX. (Jour. Bact., 36, 1938, 321.) Cells without endospores, rod-shaped, sparsely flagellated (one polar or lateral flagel- lum, or 2 to 4 peritrichous ones) ; some species non-motile. Usually Gram-negative. One genus (Chrotnobacterium) produces a violet pigment. Grow aerobically on ordi- nary culture media containing glucose. Glucose and sometimes other carbohydrates are utilized, without appreciable acid formation. Saprophytes, symbionts and pathogens. The latter are usually plant pathogens forming abnormal growths on roots and stems. Key to genera of family Rhizobiaceae. I. Cells capable of fixing free nitrogen when growing symbiotically on the roots of Leguminosae. Genus I. Rhizobium, p. 223. II. Either plant pathogens which attack roots or produce hypertrophies on stems ; or free-living non-chromogenic soil or water forms. Do not fix nitrogen. Genus II. Agrobacterium, p. 227. III. Usually free-living soil and water forms which produce a violet chromogenesis. Genus III. Chroniobacterium, p. 231. Genus I. Rhizobium Frank.* {Phytomyxa Schroeter, in Cohn, Kryptogamen-Flora von Schlesien, S, 1886, 134; Frank, Ber. d. deut. bot. Gesellsch., 7, 1889, 380; Rhizobacterium Kirchner, Beitr. z. Biol. d. Pflanzen, 7, 1895, 221; Rhizomonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 328.) From Greek rhiza, root; bios, life. Rods: 0.5-0.9 by 1.2-3.0 microns. Motile when young, commonly changing to bac- teroidal forms (a) upon artificial culture media containing alkaloids or glucosides, or in which acidity is increased; or (b) during symbiosis within the nodule. Gram- negative. Aerobic, heterotrophic, growing best with extracts of yeast, malt or other plant materials. Xitrates may be reduced to nitrites. Nitrites are not utilized. Gelatin is not liquefied or is very slightly liquefied after long incubation. Optimum temperature 25°C. This group is capable of producing nodules on the roots of Leguminosae, and of fixing free nitrogen during this symbiosis. The type species is Rhizobium leguminosarum Frank. Key to the species of genus Rhizobium. 1. Litmus milk alkaline. a. Formation of serum zone in milk. b. Moderate growth, slight acid reaction on yeast water agar plus mono-, di- and trisaccharides. c. Causes formation of root nodules on species of the genera Lathyrus, Pisum, Vicia and Lens. Bacteroids irregular with x, y, star-, and club-shaped forms; rods peritrichous when young. 1. Rhizobium leguminosarum. cc. Causes formation of root nodules on Phaseolus vulgaris, P. multiflorus and P. angustif alius . Bacteroids vacuolated rods, few branched forms ; j'oung cells peritrichous. 2. Rhizobium phaseoli. * The genus Rhizobium was revised by Dr. and Mrs. O. N. Allen under the direction of Prof. E. B. Fred and Prof. I. L. Baldwin, Univ. of Wisconsin, Madison, Wis., Jan., 1938; further revision by Dr. O. X. Allen, Jan., 1943. 224 MANUAL OF DETERMINATIVE BACTERIOLOGY ccc. Causes formation of nodules on species of genus Trifolium. Bac- teroids pear-sliaped, swollen, vacuolated. Pentoses usually not fermented. 3. Rhizobium trijolii. aa. No serum zone formed in milk. b. Scant growth, alkaline reaction on yeast water agar plus most carbohydrates. c. Causes formation of nodules on species of genus Lupinus and on Orni- thopus sativus. Bacteroids vacuolated, rods seldom branched. 4. Rhizohium lupini. cc. Causes formation of nodules on Soja max. Bacteroids long slender rods, seldom vacuolated or branched; young cells monotrichous. 5. Rhizobium japonicum.* 2. Litmus milk acid. a. Formation of serum zone in milk. b. Moderate growth, slight acid reaction on yeast w^ater agar plus mono-, di- and trisaccharides. c. Causes formation of root nodules on species of the genera Melilotus, Medicago, and Trigonella. Bacteroids club-shaped, branched, young cells peritrichous. 6. Rhizobium meliloti. 1. Rhizobium leguminosarum Frank Naturreiche. 2 Theil, Botanik, III Abt., emend. Baldwin and Fred. (Frank, Kryptogamen, Sec. 914, 1877, 1944; Landwirtschaftliche Jahrbiicher, 79, 1890, Schinzia leguminosarum Frank (all spe- 563; Rhizobium polymorphum Dangeard, cies), Bot. Ztg., 37, 1879, 377; Phyto- Rhizobiumfabae'Dangeard,'LeBotsimste, viyxa leguminosarum Schroeter (all ex- Ser. 16, 1926, 192-194 ; Baldwin and Fred, cept Rhizobium lupini), in Cohn, Krypto- Jour. Bact., 17, 1929, 146.) From Latin, gamen-Flora von Schlesien, 3, I, 1886, of the legume family (Leguminosae) . 135; Bacillus radicicola Beijerinck (all species), Bot. Ztg., 46, 1888, 726; Bacillus Note: The following binomials have fabae Beijerinck (from broad bean) and been used for species of this genus. The Bacillus oi-nilhopi Be'iierinck (from ser- names given were used by their authors radella), Bot. Ztg., 48, 1890, 837; Clado- to cover one or more of the species here chytriuin tuberculorum Vuillemin (all recognized as belonging to the genus species?), Ann. Sci. Agron. Franc, et Rhizobium. Where a question mark (?) Etrang., 5, I, 1888, 193; Bacterium is used it indicates that the species was radicicola Prazmowski (all species), too poorly described to be recognizable Landw. Vers. Sta., 37, 1890, 204; Rhizo- today. Schinzia cellulicola Frank, 1877 femmnmiobiZe Schneider (several species) (all species) Leunis, Synopsis der drei Rhizobium curvum Schneider (?), Rhizo- * No specific name has been proposed for the organism causing the formation of nodules on plants that are members of the so-called "cowpea" group. Data showing possible inter-relationships of certain plant species of the soybean and cowpea cross- inoculation groups prompted Walker and Brown (Soil Science, 39, 1935, 221-225) to propose a consolidation of the two groups to be recognized as being inoculated by a single species, Rhizobium japonicum. Results obtained recently by Reid and Bald- win (Proc. Soil Sci. Soc. Amer. for 1936, 1, 1937, 219) show these inter-relationships to include the lupine group also. FAMILY HHIZOBIACEAE 225 hium frankii var. majus and var. mi- nus Schneider (?), Rhizohium nodosum Schneider (?), Rhizohium dubium Schneider (?), Bui. Torrej' Bot. Club, 19, 1892, 213; Rhizohium sphaeroides Schneider (?), Ber. deut. bot. Gesell., 12, 1894, 16; Bacillus tuberigenus Gonner- mann and Micrococcus tuberigenus Gon- nermann, Landw. Jahrb., 33, 1894, 654, 657, are thought bj" Fred, Baldwin and McCoy (University of Wisconsin, Stud- ies in Science, Xo. 5, 1932, 140) not to be true noduJe organisms and to be too poorly described to be recognizable to- day ; Rhyzobium pasteurianum Maze (all species), Ann. Inst. Pasteur, 13, 1899, 146; Pseudorhizobium ramosnm Hartleb (?) (Chem. Zeit., U, 1900, 887) (used for noninfective culture claimed b}" Stutzer (Mitt. Landw. Inst. Breslau, /, Heft 3, 1900, 63) to be }>;enuine root nod- ule organism) ; Rhizohium radicicola Hiltner and Stormer (several species) and Rhizohium heijerinckii Hiltner and Stormer (from lupine, serradella and soy bean), Arb. Biol. Abt. f. Land-u. Forst- wirthschaft a. K. Gesundheitsamte, 3, 1903, 269; Pseudomonas radicicola ]\Ioore (all species), U. S. Dept. Agr. Bur. Plant Ind., Bui. 71, 1905, 27; Rhizomonas heijerinckii Orla-Jensen and Rhizomonas radicicola Orla-Jensen (see Hiltner and Stormer), Cent. f. Bakt., II Abt., 22, 1909, 328; Bacillus or Bacterium radici- cola Lohnis and Hansen (peritrichous species). Jour. Agr. Research, 20, 1921, 554 ; Rhizohium radicicolum Bergej' et al. , Manual, 1st ed., 1923, 40 (monotrichous species) ; Rhizohium loti Dangeard (from lotus), Rhizohium simplex Dangeard (from sainfoin), Rhizohium torulosum Dangeard (from Scotch broom), Le Botaniste, Ser. 16, 1926, 195-197. Rods: 0.5 to 0.9 by 1.2 to 3.0 microns. Motile with peritrichous flagella. Bac- teroids commonly irregular with x, y, star- and club-shaped forms. Vacuolate forms predominate. Gram-negative. Growth on mannitol agar is rapid, with tendency to spread. Streak is raised, glistening, semi-translucent, white, slimy and occasionally viscous. Considerable gum is formed. Slight acid production from glucose, galactose, mannose, lactose and maltose. Aerobic. Optimum temperature 25°C. Source : Root nodules on Lathyrus, Pisum (poa), Vicia (vetch) and Lens (lentil). Habitat : Widely distributed in soils where the above mentioned legumes are grown 2. Rhizobiumphaseoli Dangeard. (Le Botaniste, Ser. 16, 1926, 197.) From Latin, phascolus, bean; M. L. Phaseolus, a generic name. Rods : Motile with peritrichous fla- gella. Bacteroids are usuallj^ rod-shaped, often vacuolated with few branched forms. Usually smaller than in Rhizo- hium leguminosariim and R. trifolii. Gram -negative. Growth on mannitol agar is rapid with tendency to spread. Streak inoculation is raised, glistening, semi-translucent, white, slimJ^ Occasionally mucilagi- nous but this character is not so marked as in Rhizohium trifolii. Very slight acid formation from glu- cose, galactose, mannose, sucrose and lactose. Aerobic. Optimum temperature 25 ''C. Source : Root nodules of Phaseolus vulgaris (kidney bean), P. ayigustifolius (bean) and P. multiflorus (scarlet run- ner). (Burrill and Hansen, 111. Agr. Exp. Sta. Bui. 202, 1917, 137.) Habitat : Widely distributed in the soils in which beans are grown. 3. Rhizobium trifolii Dangeard. (Le Botaniste, Ser. 16, 1926, 191.) From M. L. Trifolium, a generic name. Rods : Motile with peritrichous fla- gella. Bacteroids from nodules are pear- shaped, swollen and vacuolated. Rarely X and y shapes. Gram-negative. 226 MANUAL OF DETERMINATIVE BACTERIOLOGY Growth on mannitol agar is rapid. Tlic colonies are white becoming turbid with age. Frequently mucilaginous. Streak cultures transparent at first. Growth mucilaginous later flowing down the agar slant and accumulating as a slimy mass at the bottom. Produces large amounts of gum. Slight acid production from glucose, galactose, mannose, lactose and maltose. Aerobic. Optimum temperature 25°C. Source : Root nodules of species of Trifolium (clover). Habitat: Widely distributed in the soils where clover grows. 4. Rhizobixim lupini (Schroeter) Eck- hardt, Baldwin and Fred. {Phytomyxa lupini Schroeter, in Cohn, Kryptogamen- Flora von Schlesien, 3, I, 1886, 135; Rhizobium minimum Dangeard, Le Bo- taniste, Ser. 16, 1926, 198; Eckhardt, Baldwin and Fred, Jour. Bact., 21, 1931, 273.) From Latin, Lupinus, lupine. Rods : Motile with flagella 1 to 4, usually 2 or 3. Bacteroids are vacuolate rods, seldom if ever branched. Gram-negative. Growth on yeast water, mannitol agar is scant to moderate with alkaline reaction. Beef -peptone gelatin : Little growth with extremely slow liquefaction. On galactose an alkaline reaction serves to differentiate Rhizobium lupini from all fast-growing rhizobia (R. phaseoli, R. meliloti, R. trifolii, and R. legumino- sarum). An initial alkaline reaction fol- lowed more quickly by an acid reaction on rhamnose and xylose separates R. lupini from slow-growing R. japonicxim and the Rhizobium sp. from cow pea. In general Rhizobium lupini produces slight to moderate acidity on pentose sugars and no change or alkaline reaction on hexoses, disaccharides and trisac- charides. Litmus milk : No serum zone, no re- duction, and a slight alkaline reaction. Meager growth on potato and parsnip slants, and carrot agar. Aerobic. Optimum temperature 25°C. Source : Root nodules on Lupinus (lupine), Serradella and Ornithopus. Habitat : Widely distributed in soils in which these legumes grow. 5. Rhizobium japonicum (Kirchner) Buchanan. (Rhizobacterium japonicum Kirchner, Beitrage zur Biol. d. Pflanzen, 7, 1895, 213; Pseudomonas japonica Lohnis and Hansen, Bacterium japoni- cum Lohnis and Hansen, Jour. Agr. Res., 20, 1921, 551; Rhizobium sojae Dangeard, Le Botaniste, Ser. 16, 1926, 200; Buchanan, Proc. Iowa Acad. Sci., 33, 1926, 81.) From M. L., of Japan. Rods : Motile with monotrichous fla- gella. Bacteroids of nodules are long and slender with only occasional branched and swollen forms. Gram-negative. Growth on mannitol agar is slow and scant. The streak is slightly raised, glistening, opaque, white, butyrous, with little gum formation. Pentose sugars give better growth than the hexoses. Little if any acid formed from carbo- hydrates. Acid slowly formed from xylose and arabinose. Aerobic. Optimum temperature 25°C. Source : Root nodules on Soja max (soy bean). Habitat : Widely distributed in soils where soy beans are grown. 6. Rhizobium meliloti Dangeard. (Le Botaniste, Ser. 16, 1926, 194.) From Greek, melilot, a kind of clover; M. L., Melilotus. Rods: Motile with peritrichous fla- gella. Bacteroids club-shaped and branched. Gram-negative. Growth on mannitol agar is fairly rapid. The streak is raised, glistening, opaque, pearly white, butyrous. Considerable gum is formed. FAMILY RHIZOBIACEAE 227 Acid from glucose, galactose, mannose Note : See Monograph on Root Nodule and sucrose. Bacteria and Leguminous Plants by E. Aerobic. B. Fred, I. L. Baldwin and Elizabeth Optimum temperature 2o°C. McCoy, University of Wisconsin Studies Source : Root nodules of Melilotus in Science, Madison, No. 5, 1932, xx + (sweet clover), Medicago, and Trigonella. 343 pp. for a more complete discussion of Habitat : Widelj' distributed in soils this group with an extensive bibliogra- in which these legumes grow. phy. Genus II. Agrobacterium Conn.* (.Jour. Bact., 44, 1942, 359.) From Greek, agrus, a field; M.L., bacterium, a small rod. Small, short rods which are typically motile with 1 to 4 peritrichous flagella (if only one flagellum, lateral attachment is as common as polar). Ordinarily Gram- negative. On ordinary culture media, they do not produce visible gas nor sufficient acid to be detectable by litmus. In synthetic media, enough CO2 may be produced to show acid with brom thymol blue, or sometimes with brom cresol purple. Gelatin is either very slowly liquefied or not at all. Free nitrogen cannot be fixed ; but other inorganic forms of nitrogen (nitrates or ammonium salts) can ordinarilj^ be utilized. Optimum temperature, 25° to 30°C. Habitat: Soil, or plant roots in the soil; or the stems of plants where they produce hypertrophies. The type species is Agrobacteriitm tumefaciens (Smith and Townsend) Conn. Key to the species of genus Agrobacterium. I. Plant pathogens. Produce browning of mannitol-calcium-glycerophosphate agar. Nitrate reduction weak or none. A. Nitrite produced from nitrate toaslight extent. Galls produced on plant roots. 1. Agrobacterium tumefaciens. B. Nitrite not produced from nitrate. 1. Pathogenic to apples. 2. Agrobacteriitm rhizogenes. 2. Pathogenic to raspberries and blackberries. 3. Agrobacterium rubi. II. Not pathogenic to plants. Produces browning in mannitol-calcium-glycerophos- phate agar. Nitrate reduction vigorous, with disappearance of the nitrate. 4. Agrobacterium radiobacier. 1. Agrobacterium tumefaciens (Smith 1923, 189; Conn, Jour. Bact., 44, 1942, and Town.send) Conn. (Bacterium tume- 359.) From Latin tumefaciens, swell- faciens Erw. Smith and Townsend, Sci- ing up, producing a tumor, ence, N. S. 25, 1907, 672; Pseudomonas Probable sj'nonyms : Bacillus ampelop- tumefaciens Stevens, The Fungi which sorae Trevisan, in Saccardo, S3dloge Cause Plant Disease, 1913, 35; Bacillus Fungorum, 8, 1889, 983; Bacillus am- tumefaciens Holland, Jour. Bact., 5, 1920, pelopsorae Trevisan emend. Cavara, Staz. 220; not Bacillus tumefaciens Wilson, Sperim. Agara. Ital. Modena, 30, 1897, Lancet, 1, 1919, 675; Phytomonas tume- 483; see Elliott, Bact. Plant Pathogens, faciens Bergey et al.. Manual, 1st ed., 1930, 235. * Prepared by Prof. H. J. Conn, New York State Experiment Station, Geneva, New York, September, 1943. 228 MANUAL OF DETERMINATIVE BACTERIOLOGY Among the synonyms listed in previous editions of the Manual has been Poly- monas tumefaciens Lieske, Cent. f. Bakt., I Abt., Orig., 108, 1928, 118. This is only a partial synonym, however, as its author described it as the cause of animal and human cancer, of which he regarded crown-gall of plants as merely a phase : for the origin of this theory, see Smith and Townsend, Sci., N.S. 25, 1907, 671, and Smith, Jour. Cancer Res., 7, 1922, 1-105. Description taken from the following: Riker, Banfield, Wright, Keitt and Sagen, Jour. Agr. Res., 41, 1930, 507; Sagen, Riker and Baldwin, Jour.Bact., £8, 1934, 571; Hendrickson, Baldwin and Riker, Jour. Bact., 28, 1934, 597. Rods: 0.7 to 0.8 by 2.5 to 3.0 microns, occurring singly or in pairs. Capsules. Motile with 1 to 4 flagella. Gram- negative. Agar colonies: Small, white, circular, smooth, glistening, translucent, entire. Broth: Slightly turbid, with thin pel- licle. Litmus milk: Slow coagulation. Lit- mus reduced. Neutral to alkaline. Nitrites produced from nitrates to a very slight extent. Indole: Slight amount. Slight acid from glucose, fructose, arabinose, galactose, mannitol and salicin. Starch not hydrolyzed. Optimum temperature 25° to 28°C. Facultative anaerobe. Distinctive characters : Causes a gall formation parenchymatous in character which because of its soft nature is sub- ject to injury and decay. Agrobacterium tumefaciens strongly ab- sorbs Congo red and aniline blue in con- trast to little or no absorption by A. rhizogenes. A. tumefaciens makes abun- dant growth on sodium selenite agar and calcium glycerophosphate medium with mannitol in contrast to no growth or a very slight trace by A. rhizogenes (Hen- drickson et al., Jour. Bact., 28, 1934, 597). Source: Isolated from galls on plants. Habitat : Causes galls on Paris daisy and cross-inoculable on over 40 families. 2. Agrobacterium rhizogenes (Riker et al.) Conn. (.Bacterium rhizogenes Riker, Banfield, Wright, Keitt and Sagen, Jour. Agr. Res., 41, 1930, 536; Phyto- monas rhizogenes Riker et al., ibid., 536; Pseudomonas rhizogenes Riker et al., ibid. 536; Conn, Jour. Bact., 44, 1942, 359.) From Greek, rhiza, root; genes, producing. Rods: 0.4 by 1.4 microns, occurring singly. Motile with one to 4 flagella. Encapsulated. Not acid-fast. Gram- negative. Gelatin : No liquefaction. Agar colonies : Circular, smooth, con- ve-x, finely granular; optical characters, translucent through gray to almost white. Agar slant: Moderate, filiform, trans- lucent, raised, smooth, slimy. Broth: Turbid, with heavy pellicle. Litmus milk: Acid, slow reduction. Indole not formed. Nitrites not produced from nitrates. Acid but not gas from arabinose, xy- lose, rhamnose, glucose, galactose, man- nose, maltose, lactose, salicin and ery- thritol. No acid or gas from fructose, sucrose, raffinose, melezitose, starch, dextrin, inulin, aesculin, dulcitol or man- nitol. Starch not hydrolyzed. Optimum temperature 20° to 28°C. Aerobic. Distinctive characters : Agrobacterium rhizogenes differs from Agrobacterium tumefaciens by stimulating root forma- tion instead of soft parenchymatous crown galls. A. rhizogenes lacks ability of A. tumefaciens to utilize simple nitrogenous compounds as KNO3. A. rhizogenes absorbs congo red and brom thymol blue slightly and aniline blue not at all. Will not grow on sodium selenite agar (see A. tumefaciens for response to same materials). Does not infect tomato . FAMILY RHIZOBIACEAE 229 Sources : Description made from ten cultures isolated from hairy -root of apple and other plants. Habitat: Pathogenic on apple, etc. 3. Agrobacterium rubi (Hildebrand) Starr and Weiss. {Phytomonas rubi Hildebrand, Jour. Agr. Res., 61, 1940, 694; Bacterium rubi Hildebrand, ibid., 694; Pseudomonas rubi Hildebrand, ibid., 694; Banfield, Phytopath., 20, 1930, 123; Pinckard, Jour. Agr. Res., 50, 1935, 933; Starr and Weiss, Phytopath., 33, 1943, 316.) From Latin, rubus, blackberrj' bush; M. L., Rubus, a generic name. Rods: 0.6 by 1.7 microns. Singly, in pairs or short chains. IMotile with 1 to 4 flagella. Gram-negative. Gelatin : No liquefaction. Potato-mannitol-agar slants: Growth slow, moderate, filiform, white to creamy- white, with butyrous consistency later becoming leathery. Broth: Turbid in 36 to 48 hours. Milk: A slight serum-zone, pink color, acid and curd formed. Nitrites not produced from nitrates. Ferric ammonium citrate, uric acid, oxamide, succinimide, 1-asparagine, 1- tjTosine, 1-cystine, d-glutamic acid and yeast extract can be used as a source of nitrogen (Pinckard, loc. cit.). Hydrogen sulfide not formed. Indole not formed. Acid from glucose, d-galactose, d- mannose, d-fructose, d-xylose, d-arabin- ose, sucrose, and maltose. None from lactose (Pinckard, loc. cit.). Starch not hydrolyzed. Optimum temperature 28°C. Mini- mum 8°C. and maximum 36°C. (Pinck- ard, loc. cit.). Distinctive characters. Differs from Agrobacterium tumefaciens in that it does not utilize nitrates, and grows much more slowly on ordinary media. Infects only members of the genus Rubus. Starr and Weiss (Phytopath., 33, 1943, 317) state that this species unlike Agrobac- ferivm itimefaciens and Agrobacterium rhizogcncs does not utilize asparagin as a sole source of carbon and nitrogen. Source: Isolated by Banfield {loc. cit.) and by Hildebrand {loc. cit.) from rasp- berry canes, Rubus spp. Habitat : Pathogenic on black and purple cane raspberries, and blackberries, and to a lesser extent on red raspberries. 4. Agrobacterium radiobacter (Bei- jerinck and van Delden) Conn. {Bacillus radiobacter Beijerinck and van Delden, Cent. f. Bakt., II Abt., 9, 1902, 3; Bac- terium radiobacter Lohnis, Cent, f . Bakt., II Abt., 14, 1905, 589; Rhizobium radio- bacter Pribram, Klassifikation der Schizo- myceten, Leipzig, 1933 ,5Z lAchromobacter radiobacter Bergey ct al., Manual, 4th ed., 1934, 230; Alcaligenes radiobacter Conn, in Manual, 5th ed., 1939, 97; Conn, Jour. Bact., U, 1942,359.) From Latin, mdn^s, the spoke of a wheel; Latin, bactrum, a rod. Small rods, 0.15 to 0.75 l^y 0.3 to 2.3 microns, occurring singly, in pairs and under certain conditions, in star-shaped clusters. Motile with one to four flagella. Prevailingly Gram-negative ; but an occa- sional culture is variable. Nutrient gelatin stab : No liquefaction. Agar slant: Flat, whitish slimy layer. Mannitol-calcium-glycerophosphate- agar streak plates: Abundant, raised, slimy growth surrounded by a brown halo with an outer zone of white precip- itate (Riker et al.. Jour. Agr. Res., 41, 1930, 524). Broth: Turbid; with heavy ring or pellicle if veal infusion is present. Litmus milk : Serum zone with pellicle in one week; usually turns a chocolate brown in 2 weeks; same in plain milk, but with less browning. Potato : Raised slimy mass becoming brownish; potato may be browned. Nitrates disappear (assimilated or reduced). Starch not hydrolyzed. No organic acid or visible gas from sugars; nearly all sugars, glycerol and 230 MANUAL OF DETERMINATIVE BACTERIOLOGY mannitol are utilized with the produc- tion of CO2. Optimum temperature 28°C. Mini- mum near 1°C. Maximum 45°C. Aerobic. Media containing KNO3, K2HPO4, and glycerol, ethyl or propyl alcohol become alkaline to phenol red. (Sagen, Riker and Baldwin, Jour. Bact., 28, 1934, 571.) Growth occurs in special alkaline media of pH 11.0 to 12.0 (Hofer, Jour. Amer. Soc. Agron.,^7, 1935,228). Hydrogen sulfide produced if grown in ZoBell and Feltham's medium (Jour. Bact., 28, 1934, 169). Distinctive characters : Browning of mannitol-calcium-glycerophosphate agar. Inability to cause plant disease or to produce nodules on roots of legumes. Complete utilization (disappearance of nitrate) in the peptone-salt medium of Riker et al. (Jour. Agr. Res., 41, 1930, 529) and failure to absorb congo red (ibid., 528). The species bears at least superficial resemblances to certain Rhizobium spp., but may be distinguished from them by the first two characters listed above, and the following in addition : Growth at a reaction of pH 11-12. Heavy ring or pellicle formation on veal infusion broth. H2S production in the mannitol-tryptone medium of ZoBell and Feltham (loc. cit.). Production of milky white precipitate on nitrate-glycerol-soil-extract agar. Source: Isolated from soil. Habitat: Soil, around the roots of plants, especially legumes. Note: Palacios and Bari (Proc. Indian Acad. Sci., 3, 1936, 362; Abs. in Cent. f. Bakt., II Abt., 95, 1937, 423) have de- scribed Bacillus concomitans as a sym- biont from legume nodules that has no power to fix nitrogen although it is very much like legume nodule bacteria {Rhizo- bium spp.). This organism resembles Agrobaclerium radiobacter . Appendix: The following species prob- ably belong in Agrobaclerium, but are not sufficiently well described to make their relationship certain. 1 . Agrobacterium gypsophilae (Brown) Starr and Weiss. {Bacterium gypsophilae Brown, Jour. Agr. Res., 48, 1934, 1109; Pseudomonas gypsophilae Stapp, Bot. Rev., 1, 1935, 407; Phytomonas gypso- philae Stapp, ibid., 407; Starr and Weiss, Phytopath., 33, 1943, 316.) From M. L., Gypsophila, a generic name. Rods: 0.2 to 0.8 by 0.4 to 1.4 microns. Motile with 1 to 4 flagella. Capsules. Gram-negative. Gelatin : Liquefaction slow, beginning after 1 month. Beef -infusion agar colonies : Circular, Naples yellow, smooth or rough, butyrous. Broth : Turbid in 24 hours. Milk: Coagulation and peptonization. Nitrites are produced from nitrates. Indole not produced. Hydrogen sulfide : A trace may be produced. Acid but not gas from glucose, sucrose, maltose, mannitol and glycerol. No acid from lactose. Starch not hydrolyzed. Aerobic, facultative. Distinctive characters: Differs from Xanthomonas beticola in starch hydrol- ysis, HoS production, and will not cross- inoculate with this species. Source : Isolated from several galls on Gypsophila. Habitat : Produces galls in Gypsophila panicvlata and related plants. 2. Bacterium pseudotsugae Hansen and Smith. (Hansen and R .E. Smith, Hil- gardia, 10, 1937, 576; Phytoinonas pseu- dotsugae Burkholder, in Manual, 5th ed., 1939, 209.) From M. L., Pseudotsuga, a generic name. Rods : 0.5 to 1.5 by 1.9 to 3.9 microns. Probably motile ; type of flagellation doubtful. Gram-negative. Gelatin : Liquefied. FAMILY RHIZOBIACEAE 231 Nutrient agar slant: Growth scanty, flat, glistening, smooth, translucent, whitish. Broth: Growth slight. Xo sediment. Milk: No acid. Nitrites produced from nitrates. Hydrogen sulfide production slight. Acid but not gas from glucose, fruc- tose, galactose and maltose. No acid or gas from lactose, sucrose or glycerol. Starch not hydrolyzed. Facultative aerobe. Source : Isolated from galls on Douglas fir in California. Habitat : Pathogenic on Douglas fir, Pseudoisuga taxifolia. Genus III. ChiOTD.oha.cteTiuia Bergonzini.* (Ann. Societa d. Naturalisti in Modena, Ser. 2, 14, 1881, 153.) Greek, chroma, color; M. L., bacterium, a small rod. Rods, 0.4 to 0.8 by 1.0 to 5.0 microns. Motile with 1 to 4 or more flagella. Gram- negative. A violet pigment is formed which is soluble in alcohol, but not in water or chloroform. Grow on ordinary culture media, usually forming acid from glucose, sometimes from maltose, not from lactose. Gelatin is liquefied. Indole is not pro- duced. Nitrate usually reduced to nitrite. Optimum temperature 20-25°C. but some grow well at 37°C. Usually saprophytic soil and water bacteria. The type species is Chromobacterium violaceum (Schroeter) Bergonzini. Key to the species of genus Chromobacterium. I. Motile rods. Single flagellum. A. Acid from glucose and maltose. No acid from sucrose. Nitrites produced from nitrates. No growth at 37°C. 1. Chromobacterium violaceum. II. Motile rods. Flagella generally peritrichous. A. Acid from glucose. Nitrites generallj' not produced from nitrates. Good growth at 37°C. 2. Chromobacterium ianthinum. B. Generally no acid from glucose. Nitrites produced from nitrates. No growth at 37°C. 3. Chromobacterium amethystinum. 1. Chromobacterium violaceum (Schroeter) Bergonzini. {Bacteridium violaceum Schroeter, Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 126; Micro- coccus violaceus Colin, Beitrage z. Biol, d. Pflanzen, /, Heft 2, 1872, 157; Cromo- bacterium violaceum (sic) Bergonzini, Ann. Societa d. Naturalisti in Modena, Ser. 2, 14, 1881, 153; Bacillus violaceus Schroeter, Kryptogamen-Flora von Schle- sien, 3, 1886, 157; Streptococctis violaceus Trevisan, I generi e le specie delle Batteriacee, 1889, 31 ; Pseudomonas violacea Migula, Arb. a. d. Bakt. Inst. Karlsruhe, 1, 1894, 237; Bacterium vio- laceum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 58; see 2 Aufl., 2, 1899, 262.) From Latin, violaceus, violet-colored. Note: Bacterium ianthinum Zopf (Die Spaltpilze, 1885, 68) has been regarded as identical with the above organism by Schroeter (Kryptogamen-Flora von Schlesien, 3, 1, 1886, 157), and by Leh- * Adapted by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York from Cruess-Callaghan and Gorman, Scientific Proc. Royal Dublin Society, 21, 1935, 213 in Jan. 1938; further revision, July, 1945 by Robert S. Breed with the assistance of Capt. W. C. Tobie, Sn. C, Old Greenwich, Conn. 232 MANUAL OF DETERMINATIVE BACTERIOLOGY mann and Neumann (Bakt. Diag., 1 Aufl., 2, 1896, 266; also 7 Aufl., 2, 1927, 463). Lehmann and Neumann {loc. cil.) also consider Bacillus violaceus laurenti- cus Lustig (Diagnostik der Bakterien des Wassers, 1893, 103) as being identical with Bacterium violaceum. Slender rods: 0.8 to 1.0 by 2.0 to 5.0 microns, occurring singly and in chains. Motile, with a single flagellum. Gram- negative. Gelatin colonies : Circular, gray, entire margin, assuming a violet color in the center. Gelatin stab : Infundibuliform lique- faction with violet sediment in fluid. Agar colonies : Whitish, flat, glistening, moist, becoming violet. Agar slant: Deep, violet, moist, shiny spreading growth. Broth: Slightly turbid, with violet ring and ropy sediment. Litmus milk: Violet pellicle. Diges- tion. Alkaline. Potato : Limited, dark violet growth. Loffler's blood serum : Slowly liquefied. Indole not formed. Nitrites produced from nitrates. Acid from glucose and usually from maltose. No acid from lactose or sucrose. Aerobic, facultative. Optimum, temperature 25° to 30°C. No growth at 37 °C. Slight growth at 2° to 4°C. Source : Originally grown on slices of cooked potato exposed to air contamina- tion, and incubated at room temperature. Habitat : Water. 2. Chromobacterium ianthinum (Zopf) Holland. {Bacterium ianthinum Zopf, Die Spaltpilze, 2 Aufl., 1884, 62; Bacillus janthinus Fliigge, Die Mikroorganismen, 1886, 291 ; Bacieridium ianthinum Schroc- ter, Kryptogamen Flora von Schlesien, 3, 1, 1886, 157; Pseudomonas ianthina Migula, Syst. d. Bakt., 2, 1900, 941 ; Pseu- domonas janthina Chester, Man. Determ. Bact., 1901, 317; Holland, Jour. Bact., 5, 1920, 222.) From Greek, ianthinus, violet-blue. Rods : 0.5 to 0.8 by 1.5 to 5.0 microns, occurring singly. Motile with peritri- chous flagella. Gram-negative. Gelatin colonies: Circular, yellow, becoming violet. Gelatin stab : White to violet surface growth. Infundibuliform liquefaction. Agar colonies : Creamy center, violet margin. Agar slant : Yellowish, moist, glisten- ing, becoming deep violet. Broth: Turbid, with light violet pel- licle. Litmus milk: Slow coagulation with violet cream layer. Litmus decolorized from below. Potato : Violet to violet-black, spread- ing growth. Indole not formed. Nitrites generally not produced from nitrates. Acid from glucose. No acid from maltose, lactose and sucrose. Aerobic, facultative. Optimum temperature 30°C. Grows well at 37°C. No growth at 2 to 4°C. Source : Originally grown on pieces of pig's bladder floated on badly contam inated water. Habitat : Water and soil. This may be the species that causes a fatal septi- cemia in animals and man. See Chromo- bacterium violaceum manilae. 3. Chromobacterium amethystinum (Chester) Holland. {Bacillus membra- naceus amethystinus Eisenberg, Bakt. Diag., 1891, 421; Bacterium amethystinus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 117; Bacterium membrana- ceus amethystinus Chester, ibid., 13S: Bacillus amethystinus Holland, Jour. Bact., 5, 1920, 217; not Bacillus amethys- tinus Chester, loc. cit., 262; Holland, loc. cit., 222; Bacterium memhranaceum amethystinum Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 463; Bac- teriujn violaceum amethystinum Cruess- Callaghan and Gorman, Sci. Proc. Royal Dublin Society, 21, 1935, 219.) From Greek, bluish-violet, amethyst. FAMILY RHIZOBIACEAE 233 Rods: 0.5 to 0.8 by 1.0 to l.-l microns, occurring singlj'. ^lotile with a single or occasionally with peritrichous flagella. Gram-negative. Gelatin colonies: Thin, bluish, becom- ing violet, crumpled. Gelatin stab: Heavy, violet-black pel- licle. Liquefied. Agar colonies: Deep violet, surface rugose. Agar slant: Thick, moist, yellowish- white, becoming violet with metallic luster. Broth: Pellicle with violet sediment, fluid becoming violet. Litmus milk: Violet pellicle. Diges- tion turning alkaline. Potato: Deep violet, rugose spreading growth. Indole not formed. Nitrites produced from nitrates. Usually no acid from glucose, maltose and sucrose. No acid from lactose. Aerobic, facultative. Optimum temperature 30°C. No growth at 37°C. Good growth in 7 days at 2 to 4°C. Original source : Found once by Jolles in spring water from Spalato. Habitat: Water. Appendix: The following organisms have been assigned to this genus or are believed to belong here. Additional comparative studies are badly needed. Bacillus cyaneo-fuscus Beijerinck. (Beijerinck, Bot. Ztung., .',9, 1891, 704; Bacterium ci/anojuscus Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 116 and 132.) From black glue, blue Edam cheese, water and soil. Bacillus lacyyius Schroeter. (Schroeter in Cohn, Kryptogamen-Flora von Schle- sien, 3, 1, 1889, 158.) In greenhouse on fresh paint. Bacillus lilacinus Mace. (Traite Pra- tique Bact., 6« ed., 2, 1913, 416.) From water. Bacillus tuembranuceus umethystinus inobilis Germano. (Gcrmano, Cent. f. Bakt., 12, 1892, 516; Bacillus amethysti- nus mobilis Kruse, in Fliiggc, Die Mikroorganismen, 3 Aufl., 2, 1896, 313; Bacterium amethystinus mobilis Chester, Ann. Kept. Del. Col. Agr. Exp. Sta.^ 9, 1897. 117; Bacteria?/! memhranaceus mobilis Chester, ibid.. 138.) Pseudo- monas amethyslina Migula, Syst. d. Bakt., 2, 1900, 944; Bacillus amethys- tinus Chester, Man. Determ. Bact.. 1901, 262.) From dust. Bacillus pavoninus Forster. (Forster, in van der Sleen, Sur I'examen bacterio- logique qualitatif de I'eau. Arch. Teyler, Ser. 2, Tome 4, 3 partie, 1894, No. 59, Haarlem, Heritiere Loosjes. Also see Godfrin, These, Nancy, 1934, 46.) Causes blue discoloration of Edam cheese. Bacillus polychromogoies Chamot and Thiry. (Bacille polychrome, Thiry, Compt. rend. Soe. Biol., Paris, 48, 1896, 885; Chamot and Thiry, Bot. Gaz., 30, 1900, 378.) From well water at Nancy. Probably a Pseudomonas (Tobie, per- sonal communication). Bacillus violaceus Frankland and Frankland. (Frankland and Frankland, Ztschr. f . Hyg., e, 1888, 394 ; Pseudomonas pseudoianthina IMiguIa, Syst. d. Bakt., 2, 1900, 942.) Isolated from tap water. Said to produce spores. Bacillus violaceus laurentius Jordan. (Jordan, Mass. State Bd. Health Rept., 1890. 838; Bacterium violaceus laurentius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., .9, 1897, 117; Pseudomonas laurentia Migula, Syst. d. Bakt., 2, 1900, 944; Bacillus violaceus Chester, Man. Determ. Bact., 1901, 262; Chromo- bacterium violaceum laurentium Ford, Textb. Bact., 1927, 470.) Isolated from sewage effluent. Bacillus violaceus lutetiensis Kruse. (Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 311; Bacillus lutetiensis Chester, Man. Determ. Bact., 1901, 306; Chromobacterium violaceum lutetiense Ford, Textb. Bact., 1927, 470.) From water. Bacillus violaceus sarloryi Waeldele. (These, Pharm. Strasbourg, 1938, 55.) 234 MANUAL OF DETERMINATIVE BACTERIOLOGY From dental pus. Said to form spores. Bacterium cristallino violaceum Chol- kevitch. (Cholkevitch, 1922, quoted from Godfrin, Contribution a I'etude des bacteries bleues et violettes. Th6se, Nancy, 1934, 93.) From peat. ChromobacteriimibamptoniiJieTgey et a\ . {Bacillus memhranacens arnethystinus II, Bampton, Cent. f. Bakt., I Abt., Orig., 71, 1913, 137; Bergey et al.. Manual, 1st ed., 1923, 119; Chromobacteriutn mem- branaceum amethystinum II Ford, Textb. Bact., 1927, 473.) From water. Chromobacterium coeruleum (Voges) Bergey et al. (Bacillus coeruleus Voges, Cent. f. Bakt., U, 1893, 303; Bacterium coeruleus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta.,9, 1897, 117 ; Pseudomonas coerulea Migula, Syst. d. Bakt., 2, 1900, 945; Bergey et al.. Manual, 1st ed., 1923, 120.) From water. Chromobacterium cohaerens Grimes. (Sci. Proc. Royal Dublin Society, 19, 1930,381.) From well water. Chromobacterium hibernicum Grimes. (Sci. Proc. Royal Dublin Society, 19, 1930,381.) From well water. Chromobacterium lividum (Voges) Hol- land. (Plagge and Proskauer, Zeitsch. f. Hyg., 2, 1887, 463; Bacillus lividus Voges, Cent. f. Bakt., U, 1893, 303; relationship to Bacillus lividus Zimmer- mann uncertain. Die Bakt. unserer Trink- und Nutzwasser, Chemnitz, 2, 1894, 18; Bacillus violaceus berolineiisis Kruse, in Flugge, Die Mikroorganismen, 3 Aufl., 2, 1896, 311; Bacterium lividus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 117; Bacillus berolinensis Chester, Man. Determ. Bact., 1901, 305; Holland, Jour. Bact., 5, 1920, 215.) From water. Chromobacleriuin maris-mortui Ela- zari-Volcani. (Studies on the Micro- flora of the Dead Sea, Thesis, Hebrew Univ., Jerusalem, 1940, vii and 76.) From the Dead Sea. Chromobacterium membranacemn Ber- gey et al. (Bacillus membranaceus ame- thystinus I, Bampton, Cent. f. Bakt., I Abt., Orig., 71, 1913, 135; Bergey et al., Manual, 1st ed., 1923, 119; Chromobac- terium membranaceum amethystinum I Ford, Textb. Bact., 1927, 472.) From water. Chromobacterium membranaceum ame- thystinum III Ford. (Ford, Textb. Bact., 1927, 474; Bacillus membranaceus arnethystinus III Bampton, Cent. f. Bakt., I Abt., Orig., 7i, 1913, 138.) From water. Chromobacterium membranaceum ame- thystinum IV Ford. (Ford, Textb. Bact., 1927, 474; Bacillus membranaceus arnethystinus IV Bampton, Cent. f. Bakt., I Abt., Orig., 71, 1913, 138.) From water. Chromobacterium smithii (Chester) Bergey et al. (Bacillus coeruleus Smith, Medical News, 2, 1887, 758; Bacterium coeruleus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 118; Pseudomonas smithii Chester, Man. Determ. Bact., 1901, 318; Chromobacterium coeruleum Ford, Textb. Bact., 1927, 475; not Chro- mobacterium coeruleum Bergey et al., Manual, 1st ed., 1923, 120; Bergey et al., ibid., 121.) From water. Chromobacterium violaceum manilae Ford. (Bacillus violaceus manilae Wool- ley, U. S. Dept. Int., Bur. Govt. Labs. Bull. 15, 1904 and Bull. Johns Hopkins Hosp., 16, 1905, 89; Ford, Textb. Bact., 1927, 471.) Isolated from fatal septice- mias in water buffalo (Woolley) and man (Schattenberg and Harris, Jour. Bact., U, 1942,509). More likely to be a variety of Chromobacterium ianthinum which grows at 37°C. than of C. violaceum which does not grow at 37°C. Chromobacterium viscofucatum (Harri- son and Barlow) Bergey et al. (Bacterium viscofucatum and Bacillus viscofucatus Harrison and Barlow, Cent. f. Bakt., II Abt., 15, 1905, 517; Trans. Royal Soc. Canada, 2nd Ser., 11, 1905; Bergey et al.. Manual, 1st ed., 1923, 119.) From oily butter. Probably a non-motile Pseudo- monas (Tobie, personal communication). Chromobacterium viscosum Grimes. (Cent. f. Bakt., II Abt., 72, 1927, 367.) From butter. Pseudomonas pseudoviolacea Migula. (Syst. d. Bakt., 2, 1900, 943.) From river water. FAMILY MICROCOCCACEAE 235 FAMILY V. MICROCOCCACEAE PRIBRAM.* (Jour. Bact., 18, 1929, 385.) Cells without endospores except in Sporosarcina. Cells in their free condition spherical; during division somewhat elliptical. Division in two or three planes. If the cells remain in contact after division, they are frequently flattened in the plane of last division. They occur singly, in pairs, tetrads, packets or irregular masses. Motility rare. Generally Gram-positive. Many species form a yellow, orange, pink or red pigment. Most species are preferablj^ aerobic, producing abundant growth on ordinar}^ culture media, but capable of slight anaerobic growth. A few species are strictly anaerobic. ^Metabolism heterotrophic. Carbohydrates are frequently fer- mented to acid. Gelatin is often liquefied. Facultative parasites and saprophytes. Frequently live on the skin, in skin glands or skin gland secretions of Vertcbrata. Key to the genera of family Micrococcaceae. I. Cells occur in plates, groups or in irregular packets and masses, never in chains. Pigment, when present, is yellow, orange or red. Gram-positive to Gram- negative. Genus I. Micrococcus, p. 235. II. On the animal body and in special media cells occur as tetrads. In ordinary media cells may occur in pairs and irregular masses. White to pale yellow. Genus II. Gaffkya, p. 283. III. Cells occur in regular packets. Yellow or orange pigment usually formed. Genus III. Sarcina, p. 285. Genus I. Micrococcus Cohn.* (Cohn, Beitrage z. Biol. d. Pflanzen, /, Heft 2, 1872, 153; Microsphaera Cohn, Arch. f. path. Anat., 55, 1872, 237; not Microsphaera Leveille, Ann. Sci. Nat. Bot., Ser. 3, 16, 1851, 381 ; Ascococcus Cohn, Beitrage z. Biol. d. Pflanzen, 1, Heft 3, 1875, 154; Pediococcus Balcke, Wchnschr. f. Brauerei, 1, 1884, 183; Merista Van Tieghem, Traite de Botanique, Paris, 1884, 1114; Staphylococcus Rosenbach, Mikroorganismen bei den Wundinfektions-krankheiten des Menschen, 1884, 27; Monococcns Miller, Deutsch. med. Wchnschr., 12, 1886, No. 8, 117; Botryomyccs Bollinger, Deutsch. Ztschr. f. Tiermed., 13, 1887, 77; Urococcus Miquel, Ann. Microg., /, 1888, 518; Galac- tococcus Guillebeau, Jahrb. d. Schweiz, 4, 1890, 32; Rhudococcus Zopf, Ber. d. deutsch. Bot. Gesellsch., Berlin, 9, 1891, 28; Pyococcus Ludwig, Lehrb. d. niederen Kryptog., 1892, 27; Planococcus Migula, Arb. Bakt. Inst. Karlsruhe, 1, 1894, 236; Carphococcus Hohl, Cent. f. Bakt., II Abt., 9, 1902, 338; .4Z6ococcws Winslow and Rogers, Jour. Inf. Dis., 3, 1906, 541; Aurococcus Winslow and Rogers, ibid., 540; Pedioplana Wolff,. Cent. f. Bakt., II Abt., 18, 1907,9; Melococcus Nedrigailov, Charkov Med. Zurnal, 4, 1907, 301; Solidococcus, Liquidococcus, Indolococcus and Peptonococcus Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 332; Planomerista Vuillemin, Ann. Mycol., 11, 1913, 525; Tetracoccus Orla-Jensen (in part), The Lactic Acid Bacteria, 1919, 76.) From Greek micrus, small; coccus, a grain; M. L., a sphere. Cells in plates or irregular masses (never in long chains or packets). Gram-posi- tive to Gram-negative. Growth on agar usually abundant, some species form no * The genera Micrococcus and Staphylococcus have been combined and completely revised by Prof. G. J. Hucker, New York State Experiment Station, Geneva, New York, March, 1943 so far as the aerobic species are concerned. Dr. Ivan C. Hall, Presbyterian Hospital, New York City, revised the anaerobic section, January, 1944. 236 MANUAL OF DETERMINATIVE BACTERIOLOGY pigment but others form yellow or less commonly orange, or red pigment. Glucose broth slightly acid, lactose broth generally neutral. Gelatin frequently liquefied, but not rapidly. Facultative parasites and saprophytes. The type species is Micrococcus luieus (Schroeter) Cohn. Key to the species of genus Micrococcus. 1. Aerobic to facultative anaerobic species. I. No pink or red pigment on agar media. A. Nitrites not produced from nitrates. 1. Utilize NH4H2PO4 as sole source of nitrogen.* --. - a. Yellow pigment on agar media. Not acido-proteolytic, 1. Micrococcus liiteus. aa. No pigment produced. Not acido-proteolytic. b. Utilizes urea as a sole source of nitrogen.** 2. Micrococcus ureae. bb. Does not utilize urea. 3. Micrococcus freudenreichii. aaa. Acido-proteolytic in litmus milk. 8. Micrococcus caseolyticus. 2. Do not utilize NH4H2PO4 as sole source of nitrogen. a. Yellow pigment produced. 4. Micrococcus flavus. aa. No pigment produced. 5. Micrococcus candidus. B. Nitrites produced from nitrates. 1. Utilize NH4H2PO4 as sole source of nitrogen. a. Yellow pigment on agar media. Not acido-proteolytic. b. Gelatin liquefied. 6. Micrococcus conglomeratus . bb. Gelatin not liquefied. 7. Micrococcus varians. aa. Usually not chromogenic. Actively acido-proteolytic inlitmus milk. S. Micrococcus caseolyticus. 2. Do not utilize NH4H2PO4 as sole source of nitrogen. a. Gelatin liquefied. Ferment mannitol. b. Abundant orange growth on agar media. 9a. Micrococcus pyogenes var. aureus. bb. Abundant white growth on agar media. 9b. Micrococcus pyogenes var. albus. bbb. Yellow growth on agar media. 10. Micrococcus citreus. aa. Gelatin not liquefied or very slowly liquefied. b. Abundant orange to white growth on agar media. Ferments mannitol. 11. Micrococcus aurantiacus. bb. Scant white translucent growth on agar media. Does not ferment mannitol. 12. Micrococcus epidermidis. * That is, will grow and produce acid (sometimes slowly) on slants containing 1.5 per cent washed agar, 0.1 per cent ammonium phosphate, 1.0 per cent glucose, 0.02 per cent potassium chloride, 0.02 per cent magnesium sulfate. Add brom-cresol- purple as an indicator (Hucker, N. Y. State E.xper. Sta., Tech. Bui. 100, 1924, 25; Tech. Bui. 101, 1924, 36-40) ; Manual Pure Culture Study of Bacteria. Soc. Amer. Bact., Geneva, N. Y., Leaflet II, 9th ed., 1944, 14.) FAMILY MICKOCOCCACEAE 237 II. Pink or red pigment on agar media. A. Gelatin liquefied, slowly. Produces rose-colored pigment. 13. Micrococcus roseus. B. Gelatin not liquefied. 1. Non-motile. a. Produces cinnabar-colored pigment on gelatin. 14. Micrococcus cinnabareus. aa. Produces light, flesh-colored pigment on agar slant. Ferments glycerol and mannitol. 15. Micrococcus rubens. aaa. Produces brick-colored pigment on agar slant. Does not ferment glycerol and mannitol. 16. Micrococcus rhodochrous. 2. Motile. Produces red pigment. 17. Micrococcus agilis. 2. Anaerobic species. I. Forms gas from nitrogenous media. A. Acid from glucose. 18. Micrococcus aerogenes. B. No acid from glucose. 1. No blackening of colonies in deep agar. 19. Micrococcus asaccharolyiicus. 2. Hj'drogen sulfide formed. Deep agar colonies become black. 20. Micrococcus niger. II. No gas formed from nitrogenous media. A. Acid from glucose. 1. Acid from lactose. 21. Micrococcus grigoroffi. 2. No acid from lactose. 22. Micrococcus anaerobius. 1. Micrococcus luteus (Schroeter) Cohu. {Bactcridium lutcum Schroeter, Beitr. z. Biol. d. Pflan., /, Heft 2, 1872, 119; Cohn, ibid., 153.) From Latin, luicus golden-yellow. Spheres: 1.0 to 1.2 microns, occurring in pairs and fours. Non-motile. Gram- positive. Gelatin colonies: Yellowish-white to yellow, raised, with undulate margin. Gelatin stab: No liquefaction. Agar colonies : Small, yellowish, glisten- ing, raised. Agar slant: Citron-yellow, smooth. Broth : Clear, with yellowish sediment. Litmus milk: Usually slighth- acid, not coagulated. Potato: Thin, glistening, citron-yellow growth. Indole not formed. Nitrites not produced from nitrates. Acid from glucose, sucrose and man- nitol. No acid from lactose. Starch not hydrolyzed. Ammonia produced from peptone. Utilizes NH4H2 PO4 as a source of nitrogen. Saprophytic. Aerobic. Optimum temperature 25°C. Source : Isolated by Schroeter from dust contaminations on cooked potato. Habitat : Found in skim milk and dair}^ products, and on dust particles. 2. Micrococcus ureae Cohn. (Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 1872, 158; not Micrococcus ureae Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 169; Merisla ureae Prazmowski, Biol. Cent., 8, 1888, 301; Streptococcus ureae Trevisan, I generi e le specie delle Bat- 238 MANUAL OF DETERMINATIVE BACTERIOLOGY teriacee, Milan, 1889, 31 ; Urococcus ureae Beijerinck, Cent. f. Bakt., II Abt., 7, 1901, 52; Albococcus ureae Kligler, Jour. Infect. Dis., 13, 1943, 442; Staphylococcus wreae Holland, Jour. Bact., 5, 1920, 225.) From Greek, urum, urine; M. L., urea, urea. See Micrococcus liquefaciens Migula in the appendix for references to the gelatin-liquefying form of the species. Spheres: 0.8 to 1.0 micron, occurring singly, in pairs and in clumps. Never in chains. Non-motile. Gram-variable. Gelatin colonies : Small, white, translu- cent, slimy, becoming fissured. Gelatin stab: Slight, white growth. Very slow or no liquefaction. Agar colonies: White, slightly raised. Agar slant: Grayish -white, raised, glistening, butyrous. Broth: Turbid, with viscid sediment. Litmus milk : Slightly alkaline ; litmus slowly reduced. Milk: Acid. Potato: Slight, grayish to pale olive growth. Indole not formed. Nitrites not produced from nitrates. Urea fermented to ammonium car- bonate. Acid produced from glucose, lactose, sucrose and mannitol. Starch not hydrolyzed. Ammonium salts are utilized. Ammonia produced from peptone. Saprophytic. Aerobic. Optimum temperature 25°C. Source : Isolated from fermenting urine. Habitat : Found in stale urine and in soil containing urine. Sternberg, Man. of Bact., 1893, 601; Micrococcus acidilactis Migula, Syst. d. Bakt., 2, 1900, 112; Micrococcus acidifi- cans Migula, ibid.); Micrococcus lactis viscosus Sternberg, Man. of Bact., 1893, 604; Micrococcus amarijaciens Migula, Syst. d. Bakt., 2, 1900, 100; Coccus lactis viscosi Gruber, Cent. f. Bakt., II Abt., 9, 1902, 790 {Micrococcus lactis viscosi Lohnis, Cent. f. Bakt., II Abt., 18, 1907, 144) ; Micrococcus lactis albidus Conn, Esten and Stocking, Storrs Agr. Exp. Sta. 18th Ann. Rept., 1906, 91. Spheres: 2.0 microns in diameter, oc- curing singly and in clumps, rarely in short chains. Non-motile. Gram-posi- tive. Milk gelatin colonies: Small, white, opaque. Milk gelatin stab : Inf undibuliform liquefaction. Agar colonies: White, slimy. Agar streak: White, smooth. Broth : Turbid, with white sediment. Litmus milk: Acid; coagulated; pep- tonized. Potato: Moderate white to yellow streak. Indole not formed. Nitrites not produced from nitrates. Starch not hydrolyzed. Ammonia produced from peptone. Does not utilize urea as a source of ni- trogen. Acid from glucose, lactose and sucrose. Some strains form acid from mannitol ; others from glycerol. Saprophytic. Aerobic. Optimum temperature 20°C. Habitat : Milk and dairy utensils. 3. Micrococcus freudenreichii Guille- beau. (Landwirtsch. Jahrb. d. Schweiz, 5, 1891, 135.) Named for E. v. Freuden- reich, Swiss bacteriologist. Synonyms: Micrococcus acidi lactis Krueger, Cent. f. Bakt., 7, 1890, 464 {Micrococcus acidi lactis liquefaciens Eisenberg, Bakt. Diag., 3 Aufl., 1891, 409 ; Micrococcus acidi lactici liquefaciens 4. Micrococcus flavus Trevisan. {Mi- crococcus flavus liquefaciens Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 174; Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 34; Micrococcus flavus-liquefaciens Chester, Man. De- term. Bact., 1901, 99.) From Latin, flavus, yellow. Spheres: 0.8 to 0.9 micron, occurring FAMILY MICROCOCCACEAE 239 singly, in clumps, and occasionally in fours. Occasionally cultures are found that are motile with a single flagellum. Otherwise non-motile. Gram-variable. Gelatin colonies: Small, circular, yel- lowish to yellowish-brown, somewhat serrate margin, granulated, sharply con- toured. Gelatin stab: Yellow, wrinkled surface growth with slow, crateriform lique- faction. Agar colonies: Small, pale yellowish, homogeneous, entire. Agar slant : Canary -yellow, somewhat dry, wrinkled, raised, entire. Broth : Turbid with yellowish ring and sediment. Litmus milk: Slightly acid, soft coagulum formed, with slight reduction; slowly peptonized. Potato: Slight, canary-yellow growth. Indole is not formed. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid is generally formed from glucose and lactose. Sucrose, glycerol and man- nitol generally not fermented. Ammonium salts are utilized. Ammonia produced from peptone. Non-pathogenic. Aerobic. Optimum temperature 25°C. Source : Original source not given. Habitat : Found in skin gland secre- tions, milk, dairy products, and dairy utensils. 5. Micrococcus candidus Cohn. (Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 1872, 160 ; Staphylococcus candidus Holland, Jour. Bact., 5, 1920, 223.) From Latin candidus, shining white. Spheres: 0.5 to 0.7 micron, occurring singly. Non-motile. Gram-positive. Gelatin colonies : White, granular, with irregular or entire margin. Gelatin stab: White surface growth. Filiform. No liquefaction. Agar colonies: Punctiform, white, smooth, entire, iridescent. Agar slant: Smooth, white, glistening, iridescent. Broth: Turbid, with pellicle. Litmus milk : Slightly acid ; not coagu- lated. Potato: Thick, porcelain white, glistening. Indole not produced. Nitrites not produced from nitrates. Starch not hydrolyzed. Ammonia produced from peptone. Ammonium salts not utilized. Acid from glucose, sucrose, lactose and glycerol. Non-pathogenic. Aerobic. Optimum temperature 25°C. Source : Originally appeared as white colonies on cooked potato exposed to dust contaminations. Habitat: Found in skin secretions, milk and dairy products. 6. Micrococcus conglomeratus Migula. (Citronengelber Diplococcus, Bumm, Der Mikroorganismen der gonorrhoischen Schleimhauterkrankungen, 1 Aufl., 1S85, 17; Micrococcus citreus conglomeratus Flligge, Die Mikroorganismen, 2 Aufl., 1886, 182; Diplococcus citreus conglomer- atus Bumm, ibid., 2 Aufl., 1887; Neisseria citrea TrevLsan, I generi e le specie delle Batteriacee, Milan, 1889, 32; Mcrismo- pcdia citreus conglomeratus Dyar, Ann, N. Y. Acad. Sci., 8, 1895, 352; Migula, Syst. d. Bakt., 2, 1900, 146; not Micro- coccus conglomeratus Weichselbaum, 1887, see Trevisan, loc. cit., 33; Micrococcus citreus Winslow and Winslow, Systematic Relationships of the Coccaceae, 1908, 218.) From Latin, conglomeratus, rolled together, crowded. Spheres: 0.8 to 1.2 microns, occurring singly, in pairs, in fours, and in large clumps. Non-motile. Gram-variable. Gelatin colonies: Small, circular, yel- low with radiate margin. Gelatin stab : Slow crateriform lique- faction. 240 MANUAL OF DETERMINATIVE BACTERIOLOGY Agar colonies : Luxuriant, moist, sulfur yellow. Agar slant: Light yellow, plumose, slightly rugose, somewhat dull, raised center and transparent margin. Broth: Turbid, with light orange ring and sediment. Milk: Generally acid but not suffi- cient to curdle. Potato: No growth. Indole not formed. Nitrites produced from nitrates. Blood not hemolyzed. Starch not hydrolyzed. Acid from glucose and lactose gen- erally, sometimes from sucrose. Manni- tol and glycerol generally not fermented. Ammonia produced from peptone. Utilizes NH4H2PO4 as a source of nitrogen. Resistant to drying and heat. Non-pathogenic. Aerobic . Optimum temperature 25°C. Source : Found in gonorrhoeal pus and dust. Habitat: Infections, milk, dairy prod- ucts, dairy utensils, water, common. 7. Micrococcus varians Migula. (Merismopedia Jlava varians Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 346; Migula, Syst. d. Bakt., 2, 1900, Ido; Merismopedia flava-varians Chester, Man. Determ. Bact., 1901, 103; Micrococcus lactis vari- ans Conn, Esten and Stocking, Storrs Agr. Exp. Sta. Rept. for 1906, 121.) From Latin, varians varying. Spheres: 0.8 to 1.0 micron, occurring singly, in pairs and in fours. Occa- sionally cultures are found that are motile with a single flagellum. Otherwise non- motile. Gram -variable. Gelatin colonies: Small, circular, whit- ish to yellow, capitate, moruloid. Gelatin stab : Scant growth. No lique- faction. Agar colonies: Small, yellow, raised, glistening. Agar slant: Plumose, yellow, vari- egated. Broth: Turbid, with yellow, granular sediment. Litmus milk : Acid; coagulated on boil- ing. Potato: Raised, dry, bright-yellow, glistening. Indole not formed. Nitrites produced from nitrates. Acid from glucose, lactose, sucrose, raffinose and frequently from glycerol and mannitol. No acid from salicin or inulin. Starch not hydrolyzed. Ammonia produced from peptone. Utilizes NH4H2PO4 as a source of nitrogen. Saprophytic. Aerobic. Optimum temperature 25°C. Source : Original strains found in a contaminated jar of sterilized milk. Habitat : Has been found in body secretions, dairy products, dairy utensils, dust and water, including sea water. 8. Micrococcus caseolyticus Evans. (Evans, Jour. Inf. Dis., 18, 1916, 455; Micrococcus casei Hucker, N. Y. Agr. Exp. Sta. Tech. Bull. 102, 1924, 17; Prob- ably Micrococcus casei Holland, Jour. Bact., 5, 1920, 223.) Identical in part with Micrococcus casei acidoproteolyticus I and // Gorini, Rev. Gen. du Lait, 8, 1910, 337; Tetracoccus liquejaciens Orla-Jensen, The Lactic Acid Bacteria, 1919, 80 {Micrococcus casei liquefaciens Orla-Jensen, Doktordispu- tats, 1904; Tetracoccus casei liquefaciens Orla-Jensen, The Lactic Acid Bacteria, 1919, 80 ; Micrococcus liquefaciens Holland , Jour. Bact., 5, 1920, 224. Also see ref- erences under Streptococcus liquefaciens.) From Latin, caseus, cheese, casein; and Greek, lyticus, able to dissolve; M. L., dissolving, digesting. Spheres, variable in size, occurring in clumps. Non-motile. Gram-positive. Gelatin stab: Liquefaction generally FAMILY MICROOOCCACEAE 241 begins after first day and continues rapidly. Agar colonies : Yellow to orange (Evans, loc. cit.), pearly white (Hucker, loc. cit.). Agar stroke : Yellow to orange (Evans, loc. cit.), pearly white (Hucker, ?oc. cii.), luxuriant growth. Broth : Generally grows with smooth turbidity although certain strains give heavy precipitate with clear supernatant fluid. Litmus milk: Acid, peptonized. Whej- generally clear. Potato : Scanty white growth. Certain strains may show yellow pigment. Indole not formed. Nitrites usually produced from ni- trates. Acid from glucose, lactose, maltose, mannitol and glycerol. Xo action on raffinose. Forms dextrorotarj' lactic acid (Orla- .Tensen, 1919, loc. cit.). Asparagin and urea decomposed by some strains. Utilizes NH4H2PO4 as a source of nitro- gen. Optimum temperature 22°C. Aerobic. Saprophytic. Source : Eight cultures from bovine udder. Habitat: Milk and dairy products, especially cheese, dairy utensils. 9a. Micrococcus pyogenes var. aureus (Rosenbach) Zopf . (Staphylococcus pyo- genes aureus Rosenbach, Mikroorganis- men bei den Wundinfectionskrankheiten des Menschen, Wiesbaden, 1884, 19; Staphylococcus aureus Rosenbach, ibid., 27; Micrococcus pyogenes var. aureus Zopf, Die Spaltpilze, 3 Aufl.. 1885, 56; Micrococcus aureus Zopf, ibid., 57; Micrococcus pyogenes Lehmann and Xeu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 165; Aiirococcus aureus Winslow and Rogers, .Jour. Inf. Dis., 3, 1906, 554; Micrococcus lactis varians Conn, Esten and Stocking, Storrs Agr. Exp. Sta. Rept. for 1906, 121 ; Staphylococcus pyogenes Andre wes and Gordon, Rept. (35th) Med. Officer Local Govt. Board, London, 1907, 549; (Tetracoccus) Micrococcus pyogenes aureus Orla-Jensen, The Lactic Acid Bacteria, 1919, 81 ; Staphylococcus pyo- genes-aureus Holland, Jour. Bact., 5, 1920, 225.) From Greek, pyon, pus; M. h., -genes, producing. From Latin, aureus, golden. Spheres: 0.8 to 1.0 micron, occurring singl}', in pairs, in short chains, and in irregular clumps. Xon-motile. Gram- positive. Gelatin stab : Saccate liquefaction with yellowish pellicle and yellow to orange sediment. Agar colonies: Circular, smooth, yel- lowish to orange, glistening, butyrous, entire. Agar slant : Abundant, opaque, smooth, flat, moist, yellowish to orange. Broth : Turbid with yellowish ring and sediment, becoming clear. Litmus milk: Acid; coagulated. Potato: Abundant, orange, glistening. Indole not formed. Nitrites produced from nitrates. Acid from glucose, lactose, sucrose, mannitol and glycerol, but not from raffinose, salicin or inulin. Forms inactive or levorotary lactic acid (Orla-Jensen, loc. cit.). Slight H2S formation. Starch not hydrolyzed. Does not utilize XH4H;P04, as a source of nitrogen. Ammonia produced from peptone. Pathogenic. Individual strains vary in their ability to produce hemolysin, coagulase and other metabolic products. Certain strains, under favorable con- ditions, produce not only exotoxins (hematoxin, dermatoxin, lethal toxin, etc.) but also a potent enterotoxin which is a significant cause of food poi- soning (Dolman and Wilson, Jour. Immunology, 35, 1938. 13). Aerobic, facultative. Optimum temperature 37°C. Source : Isolated from pus in wounds. 242 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : Skin and mucous membranes. The cause of boils, abscesses, furuncles suppuration in wounds, etc. 9b. Micrococcus pyogenes var. albus (Rosenbach) Schroeter. {Staphylococcus pyogenes albus Rosenbach, Mikroorganis- men bei den Wundinfektionskrankheiten des Menschcn, Wiesbaden, 1884, 2 ; Staph- ylococcus albus Rosenbach, ibid., 27; Micrococcus pyogenes var. albus Schroe- ter, in Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 147; Micrococcus pyogenes Migula, Syst. d. Bakt., 2, 1900, 87; Alba- coccus pyogenes Winslow and Rogers, Jour. Inf. Dis., 3, 1906, 544; Micrococcus oZ6ws Buchanan, Veterinary Bacteriology 1911, 196; (Tetracoccus) Microccocus pyogenes albus Orla-Jensen, The Lactic Acid Bacteria, 1919, 81; Staphylococcus pyogenes-albus Holland, Jour. Bact., 5, 1920, 225.) From Latin, albus, white. Spheres: 0.6 to 0.8 micron, occurring singly, in pairs and in irregular groups. Non-motile. Gram-positive. Gelatin stab : Saccate liquefaction with heavy white sediment. Agar colonies : Circular, white, smooth, glistening, entire. Ten per cent evaporated milk agar: Growth at 20°C frequently orange (Chapman, Jour. Bact., 45, 1943, 405). Agar slant: Abundant, white, smooth, glistening. Broth: Turbid, with delicate pellicle and white sediment. Litmus milk : Acid ; coagulated. Little or no visible peptonization. Potato: Thick, smooth, white, glisten- ing. Indole not formed. Nitrites produced from nitrates. Hydrogen sulfide is formed. Acid formed from glucose, lactose, sucrose, glycerol and mannitol, but not from raffinose, salicin and inulin. Forms inactive or levorotary lactic acid (Orla-Jensen, loc. cit.). Starch not hydrolyzed. Ammonia produced from peptone. Does not utilize NH4H2PO4 as a source of nitrogen. Pathogenic. Production of toxins, coagulase and hemolysin as in Micro- coccus aureus. Aerobic, facultative. Optimum temperature 37°C. Source : Originally isolated from pus. Habitat : Skin and mucous membranes. Occurs in wounds, boils, abscesses, etc. 10. Micrococcus citreus Migula. (Staphylococcus pyogenes citreus Passet, Aetiologie der eiterigen phlegmone des ]\Ienschen, Berlin, 1885, 9; Micrococcus pyogenes citreus Schroeter, in Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 147; Migula, Syst. d. Bakt., 2, 1900, 147; Staphylococcus citreus Bergey et al., Manual, 1st ed., 1923,55.) From Latin, citreus, of or relating to the citrus tree; M. L., lemon yellow. Spheres: 0.9 micron, occurring singly. Gram-positive. Gelatin colonies : Circular, pale yellow, granular, entire, liquefying in 6 days. Gelatin stab : Lemon yellow surface growth sinking into the medium. Gray- ish-white growth in stab. Complete liquefaction in 43 days. Agar colonies: Small, yellow, smooth, entire. Agar slant: Broad, lemon yellow, glis- tening, elastic. Broth: Turbid, with yellow sediment and pellicle. Litmus milk: Acid, with slow coagula- tion. Potato : Thin, grayish streak, becoming citron yellow. Indole not formed. Nitrites produced from nitrates. Starch not hydrolyzed. Acid from glucose, lactose, sucrose, raffinose, inulin, salicin, glycerol and mannitol. Does not utilize NH4H2PO4 as a source of nitrogen. Ammonia produced from peptone. Aerobic, facultative. Pathogenic. Optimum temperature 37 °C. Source : Originally isolated from pus. FAMILY MICROCOCCACEAE 243 Habitat : Skin and mucous membranes of vertebrates. 11. Micrococcus aurantiacus (Schroe- ter) Cohn. (Bacteridium aurantiacuin Schroeter, Beitr. z. Biol., 1, Heft 2, 1872, 126; Colin, Beitr. z. Biol., 7, Heft 2, 1872, 154; Pediococcus aurantiacus De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1051; Micrococcus aurantiacus- sorghi Bruyning, Arch. Neer. Sci. E.xaet. et Nat., 1, 1898, 297; Streptococcus aurantiacus Chester, Man. Determ. Bact., 1901, 69; Aurococcus aurantiacus Winslow and Winslow, Systematic Rela- tionships of the Coccaceae, 1908, 186; Sarcina aurantiaca Holland, Jour. Bact., 5, 1920, 225 (not Sarcina aurantiaca Flijgge, Die Mikroorganismen, 2 Aufl., 1886, 180); Staphylococcus aurantiacus Holland, ibid.) From Latin, aurum, gold;M. L., aurantium , the orange ;M.L., aurantiacus, orange-colored. Spheres: Slightly ellipsoidal, 1.3 to 1.5 microns, occurring singly, in short chains and in small clumps. Xon-motile. Gram-positive. Gelatin colonies: Circular to oval, smooth, glistening with yellow to orange center. Gelatin stab : Yellow surface growth . No liquefaction. Agar colonies: Circular, smooth, glis- tening, yellow to orange, entire. Agar slant : Buff to scant orange-j^ellow, beaded growth, raised, glistening. Broth : Turbid, with pellicle. Litmus milk: Faintly acid, no coagu- lation. Potato : Slimy, yellow growth. Pig- ment is insoluble in alcohol and ether. Indole not produced. Nitrites generally produced from ni- trates. Slight acidity from glucose, fructose, sucrose, lactose and mannitol. Xo acid from raffinose, salicin, inulin. Starch not hydrolyzed. Ammonia produced from peptone. No growth in ammonium media. May be pathogenic. Optimum temperature 25°C. Aerobic. Source : First isolated from colonies that grew on boiled egg exposed to dust contamination. Habitat : Usually isolated from infec- tions but also found in milk, cheese and dust. Note: Albococcus cpidermidis (var. A) Kligler (Jour. Infect. Dis., 12, 1913, 444) which was based on a white culture received from Krai under the name Micrococcus aurantiacus was apparently a white strain of this organism as it grew luxurianth^ on ordinary agar. 12. Micrococcus epidermidis (Winslow and Winslow) Hucker. [Staphylococcus epidermidis albus Welch, Amer. Jour, of Med. Sci., Phila., X. S., 102, 1891, 441; Micrococcus epidermidis albus Randolph, Jour. Amer. Med. Assoc, 31, 1898, 706; Albococcus epidermidis Winslow and Win- slow, Syst. Relationships Coccaceae, New York, 1908, 201; Staphylococcus epidermidis Evans, Jour. Inf. Dis., 15, 1916, 449; Hucker, X. Y. Agr. Exp. Sta. Tech. Bull. 102, 1924, 21.) From Greek, epidermis, the outer skin. Spheres: 0.5 to 0.6 micron, occurring singly, in pairs and in irregular groups. Xon-motile. Gram-positive. Gelatin stab : White surface growth with slow saccate liquefaction. Agar colonies: Rather scant, white, translucent. Broth: Turbid, with white ring and sediment. Litmus milk: Acid. Potato : Limited growth, white. Indole not formed. Xitrites are produced from nitrates. Usually does not utilize XH4H2PO4 as a source of nitrogen. Acid formed from glucose, fructose, maltose, lactose and sucrose, but not from mannitol, raffinose, salicin or inulin. Usually fails to hemolyze blood. Xo coagulase produced. Parasitic rather than pathogenic. Aerobic, facultative. 244 MANUAL OF DETERMINATIVE BACTERIOLOGY Optimumtemperature 37°C. Source: Originally isolated from small stitch abscesses and other skin wounds. Habitat : Skin and mucous membranes. 13. Micrococcus roseus Fliigge. (Rosafarbiger Diplococcus, Bumm, Der Mikroorganismen der gonorrhoischen Schleimhauterkrankungen, 1 Aufl., 1885, 25; Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 183; Neisseria roseaTrevisan, I generi e le specie delle Batteriacee, Milan, 1889, 32; Diplococcus roseus Eisen- berg,Bakt.Diag.,3Aufl.,1891,12;MeWs- mopedia rosea Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 354; Rhodococcus roseus Winslow and Rogers, Jour. Inf. Dis., S, 1906, 545.) From Latin, roseus, rose- colored. Spheres: 1.0 to 1.5 microns, occurring singly and in pairs. Non-motile. Gram- variable. Gelatin colonies : Rose surface growth usually with slow liquefaction. Agar colonies: Circular, entire, rose- red surface colonies. Agar slant: Thick, rose-red, smooth, glistening streak. Broth : Slightly turbid with rose- colored sediment. Litmus milk: Unchanged to alkaline, usually reddish sediment after 14 days. Usually produce nitrites from nitrates. Potato: Raised, rose-red, smooth, glis- tening. Starch not hydrolyzed. Acid from glycerol and mannitol. Utilizes NH4H2PO4 as a source of nitrogen. Saprophytic. Aerobic. Optimum temperature 25°C. Source: Dust contamination. Habitat: Widespread, as it occurs in dust. 14. Micrococcus cinnabar eus Fliigge. (Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 174; Rhodococcus cinnabareus Wins- low and Rogers, Jour. Inf. Dis., 3, 1906, 545.) From M. L., cinnabar-colored. Spheres: 1.0 micron, occurring singly and in pairs. Non-motile. Gram-vari- able. Gelatin colonies : Small, circular, bright red, becoming cinnabar red. Gelatin stab : Thick, raised, rose to cinnabar red growth on surface. No liquefaction. White colonies along stab. Agar slant: A carmine-red streak. Slow growth. Broth: Turbid. Litmus milk : Slightly alkaline to slightly acid. Potato : Slowly developing vermillion red streak. Small amount of acid from test sugars. Indole not formed. Does not utilize NH4H2PO4 as a source of nitrogen. Nitrites produced from nitrates. Starch not hydrolyzed. Saprophytic. Aerobic. Optimum temperature 25°C. Source : Found as contamination of cultures. Habitat : Usually found as a dust con- tamination. 15. Micrococcus rubens Migula. {Mi- crococcus tetragenus ruber Bujwid, in Schneider, Arb. bakt. Inst. Karlsruhe, 1, Heft 2, 1894, 215; Migula, Syst. d. Bakt., ^, 1900, 177; Micrococcus ruber and Rhodococcus ruber Holland, Jour. Bact., 6, 1920, 224; Micrococcus roseofulvus Hucker, N. Y. S. Agr. Exp. Sta. Tech. Bull. 135, 1928, 27; not Micrococcus roseo- fulvus Lehmann and Neumann, Bakt. Diag., 1 Aufl., ^, 1896, 177 and 439 ; Rhodo- coccus roseoftdvus Pribram, Klassifika- tion der Schizomyceten, 1933,44). From Latin, rubens, ruddy. The following description is taken from Migula {loc. cit.) and from Hucker {loc. cit.) supplemented from unpublished notes of the latter. Also see Breed (Jour. Bact., 45, 1943,455). Spheres: 1.3 to 4.0 microns, average size 2.1 microns, occurring in fours and FAMILY MICROCOCCACEAE 245 in irregular masses, generally not singly or in pairs. Non-motile. Gram-nega- tive to Gram-variable Gelatin colonies : After several days, small, pink or flesh-colored, shiny, buty- rous, 0.5 to several mm. in diameter. Smaller colonies have regular edges ; larger colonies have lobate edges. Gelatin streak: Thick, shiny, flesh- colored to carmine-red growth, generally spreading. Gelatin stab: Scant, whitish growth along line of stab; surface growth flesh- red. No liquefaction after several weeks, but a slight softening of the medium underneath the growth. Agar slant: Luxuriant, thick, spread- ing, slimy, flesh-colored growth. Broth: Bright red, slimy sediment. No pellicle. Milk: General!}' acid curd followed by slight peptonization. Nitrites produced from nitrates. Acid from glucose, sucrose, mannitol and glycerol. No action on lactose or starch. Pigment soluble in ether, benzol, car- bon bisulfide, chloroform and alcohol. Not soluble in water (Schneider, loc. cit.). Saprophytic. Grows well at 26" to .37°C. Aerobic. Source : Original culture isolated by Bujwid in Bern, Switzerland and sent to Migula at Karlsruhe, Germany. Habitat: Unknown. 16. Micrococcus rhodochrous (Zopf) -Migula. (Rhodococcus rhodochrous Zopf, Berichte d. deutsch. bot. Gesellsch., 9, 1891, 22; Migula, Syst. d. Bakt., 3, 1900, 162.) From Greek, rhodiim, rose; chros, color. Spheres: 0.5 to 1.0 micron, occurring singly. Non-motile. Gram-variable. Gelatin colonies: Small, circular, glis- tening, raised, entire, dark, reddish- brown . Gelatin stab: Dark, carmine-red, dry surface growth. Slight growth in stab. No liquefaction. Agar slant : Carmine-red streak, be- coming brick-red in color. Broth : Thick rose-red pellicle with red, flocculent sediment. Litmus milk : Slightly alkaline. Potato: Carmine-red streak. Does not ferment glycerol and man- nitol. Aerobic. Saprophytic. Optimum temperature 25°C. Habitat : Water. 17. Micrococcus agilis Ali-Cohen. (Ali-Cohen, Cent. f. Bakt., 6, 1889, 36; Planosarcina agilis Migula, in Engler and Prantl, Die naturl. Pflanzenfam., 1, la, 1895, 20; Micrococcus agilis ruber Peppier, Cent. f. Bakt., I Abt., 29, 1901, 352; Planococcus agilis Chester, Man. Determ. Bact., 1901, 115; Rhodococcus agilis Winslow and Rogers, .Jour. Inf. Dis., 3, 1906, 545; Sarcina agilis Ender- lein, Sitzber. Gesell. Naturf. Freunde Berlin, 1930, 182; not Sarcina agilis Matzuschita, Zeit. f. Hyg., 35, 1900, 496; not Sarcina agilis Saito, Jour. Coll. Sci. Imp. Univ. Tokyo, 23, 1908, .) From Latin, agilis, agile. Spheres, 1.0 micron, occurring singly, in pairs and in fours. Motile by means of one or two flagella. Gram-variable. Gelatin colonies : Small, gray, becoming distinctly rose-colored. Gelatin stab: Thin, whitish growth in stab. On surface thick, rose-red, glis- tening growth. Generally no liquefac- tion . Agar slant: Glistening, dark rose-red, lobed, much variation in color. Broth: Slightly turbid, with slight, rose-colored ring and pink sediment. Litmus milk: Slightly acid, pink sedi- ment . Potato: Slow growth as small, rose- colored colonies. Loeflier's blood serum: Pink, spread- 246 MANUAL OF DETERMINATIVE BACTERIOLOGY ing, shiny, abuudaut. Slow liquefaction. Indole not formed. Nitrites produced (trace). Ammonia formed (trace). Does not utilize NH4H2PO4 as source of nitrogen. Acid from glucose, sucrose, inulin, glycerol and mannitol. No acid from raffinose . Aerobic. Saprophytic. Optimum temperature 25°C. Source : Isolated from water. Habitat : Water, sea water, on sea fish. *18. Micrococcus aerogenes (Schott- miiller) Bergey et al. {StapMjlococcus aerogenes Schottmliller, Cent. f. Bakt., I Abt., Orig., 64, 1912, 270; Bergey et al., Manual, 1st ed., 1923, 70; not Micrococcus aerogenes Miller, Deutsch. mod. Wchnschr., 12, 1886, 119.) From Greek, forming air or gas. Description according to Prevot, Ann. Sci. Nat., Ser. Bot. et Zool., 15, 1933, 212. Spheres: 0.6 to 0.8 micron, occurring in clusters, sometimes in pairs or short chains. Gram-positive. Gelatin: No liquefaction. Deep agar colonies: Small, lenticular, nearly spherical, yellowish white. Some gas bubbles produced, not fetid. Blood agar colonies: Very small, gray- ish. No true hemolysis, but a narrow clear zone is formed. Serum agar: Colonies lenticular. Gas not fetid. Neutral red serum agar : Colonies len- ticular. Gas produced. Neutral red changed to greenish yellow. Glucose broth with blood: Turbid. Gas produced. Hydrogen sulfide not produced. Slight hemolysis. Glucose serum broth: Turbid. Gas produced. Peptone water with serum : Gas . Indole produced. Milk: Growth feeble. Neither acid nor coagulated. Proteins not attacked. Glucose and fructose attacked slightly by two out of three strains. Does not plasmolyse readily. Neutral red broth : Changed to yellow- ish green. Nitrites not produced from nitrates. Optimum pH 6.5 to 8.0. Optimum temperature 37°C. Pathogenic. Strict anaerobe. Distinctive character: Fermentation of glucose and gas production from peptones. Source : Isolated (Schottmliller) from cases of puerperal fever. Three strains from infected tonsils studied by Prevot. Habitat : Natural cavities, especially the tonsils and female genital organs. 19. Micrococcus asaccharolyticus (Dis- taso) comb. nov. {Staphylococcus asac- charolyticus Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 445.) From Greek, not dissolving sugar. Description according to Prevot, Ann. Sci. Nat., Ser. Bot., 15, 1933, 211. Large spheres: 1.0 to 1.2 microns, occurring in very large clusters, also in pairs and short chains. Gram-positive. Gelatin: At 37°C, growth resembles tufts of cotton wdiich precipitate. No liquefaction. Deep agar colonies: Very delicate, pin-point, transparent. A few bubbles of gas produced. Broth: Turbid. Growth settles at the bottom of the tube as a sort of vis- cous zooglea. Unpleasant odor pro- duced. Peptone water: Turbid. Indole pro- duced. Milk: Feebly acidified, but not coagulated. Egg white not attacked. Carbohydrates not attacked. Strict anaerobe. Anaerobic section revised by Dr. Ivan C. Hall, New York, N. Y. FAMILY MICROCOCCACEAE 247 Distinctive characters: Large size; unpleasant odor; production of indole; production of gas. Source: Isolated from the large in- testine of a man with intestinal in- toxication. Habitat: Intestine. Not common. Note: Weinberg, Nativclle and Prevot (Les Microbes Anaerobies, 1937, 1023) regard Micrococcus indolicus Christian- sen (Ac. Pat. Micr. Scand., 18, 1934, 42) as a variety of this species giving it the name Staphylococcus asaccharolyticus var. indolicus. This variety differs from the species by forming opaque lens-shaped colonies and by a more abundant production of gas from peptone. 20. Micrococcus niger Hall. (Jour. Bact., ^0, 1930, 409.) From Latin, niger, black. Small spheres: 0.6 micron in diameter, occurring in irregular masses, occasionallj- in pairs. Gram-positive. Gelatin: After 5 days a dark sediment is produced which gradually gets more; and more intensely black. No lique- faction. Deep agar colonies: Slow growth. At first very tiny, colorless, irregularly globular, smooth, dense. Small bubbles of gas sometimes produced. After sev- eral days colonies become brown, then black. If exposed to air, colonies fade to a dull gray. Medium not discolored. Blood agar slant: After 4 or 5 days, minute, black colonies, round, smooth, glistening, 0.5 mm. in diameter. Non- hemolytic. Broth : After 4 or 5 days uniform tur- bidity and slight production of gas which contains H2S. Black sediment. Coagulated serum: Minute, brown colonies appear on the 8th day. No liquefaction. Milk: No change. Brain medium : Turbid after 4 or 5 days at 37°C. Uniform gas production about the Gth day. Discoloration of tiie me- dium not marked. No acid from carbohydrates. Black sediment produced. Non-pathogenic for guinea-i)igs and rabbits. Optimum temperature 37°C. No growth below 30°C. Strict anaerobe. Distinctive characters : Formation of a water-insoluble, black pigment. Growth slow, visible after 2 to 4 days. Source: Isolated from urine of an aged woman. Habitat: Unknown. 21. Micrococcus grigorofii Prevot. (Micrococcus A, Grigoroif, These de Geneve, 1905; Prevot, Ann. Sci. Nat., Ser. Bot. et Zool., IS, 1933, 219.) Named for Grigoroff, who first isolated this organism. Small spheres : Average size 0.7 micron, occurring singly or in irregular masses. Gram-positive. Gelatin : Colonies appear in four days. No liquefaction. Deep agar colonies: After three days, round, lenticular, yellowish. Glucose broth : Turbid after 2 days with whitish sediment. Neither gas nor fetid odor produced. The medium is acidified. Milk: Good growth. Acid. Coagu- lation. Acid from glucose, maltose, lactose, fructose and sorbitol. One strain slightly pathogenic. Optimum temperatvu'e 37°C. Strict anaerobe. Distinctive characters : This is the only anaerobic coccus growing in irregular masses that coagulates milk. Lactose is fermented. Source: Five strains isolated from the appendix by Grigoroff. One strain iso- lated from an appendix by Prevot. Habitat : Human digestive tract. Not common. 22. Micrococcus anaerobius (Hamm) comb. 710V. ("Anaerobic staphylococcus, Jungano, Compt. rend. Soc. Biol. Paris, 59, 1907, 707; Sta'phylococcus anaerobius 248 MANUAL OF DETERMINATIVE BACTERIOLOGY Hamm, Die puerperale Wundinfektion, Berlin, 1912; not Staphylococcus anaero- bius Heurlin, Bakt. Unters. d. Keimge- haltes im Genitalkanale d. fiebernden Wochnerinnen, Helsingfors, 1910, 120.) From Greek, living without air. Description according to Pr^vot, Ann. Sci. Nat., S^r. Bot., 15, 1933, 209. Small spheres: 0.5 to 0.6 micron, occurring in masses. Gram-positive. Gelatin: No liquefaction. Deep agar colonies: Lenticular, thick. No gas produced. Broth: Turbid, later clearing. Sedi- ment. Glucose broth: Good growth. Nei- ther acid nor gas produced. Peptone water: No turbidity. No gas. Indole not produced. Milk: Neither coagulated nor acidified. Coagulated serum not attacked. Egg white not attacked. Carbohydrates not attacked by the strains of Jungano. Acid feebly pro- duced from glucose and galactose by Prevot's strain. Does not plasmolyse. Temperature relations: Optimum 36° to 38°C. At 22°C growth slow, poor. No growth below 22°C. Killed in ten minutes at 80°C or in half an hour at 60°C. Optimum pH 6.0 to 8.0. Pathogenic for guinea-pigs and rabbits. Strict anaerobe. Distinctive characters: Neutral red broth remains unchanged. No gas produced. Source: First isolated by Jungano from a case of cystitis . Found by Pr^vot in the pus from a suppurated tonsil. Habitat: Urinary tract, urethra, in- testine, buccal cavity and conjunctiva. Appendix I*: The following genus is organized ona physiological basis. Because of this no attempt is made to fit it into the classification outline. A single specieshas been described. Genus A. Methanococcus Kluyver and van Niel. (Cent. f. Bakt., II Abt., 94, 1936, 400.) Spherical cells, occurring singly or in masses. Motility not observed. No endo- spores formed. Gram-variable. Chemo-heterotrophic, anaerobic, fermenting vari- ous organic compounds with the formation of methane. Saprophytes. The type species is Methanococcus mazei Barker. 1. Methanococcus mazei Barker. (Pseudosarcina, Maze, Compt. rend. Soc. Biol., Paris, 78, 1915, 398; Barker, Arch, f . Mikrobiol., 7, 1936, 430.) Named for Maze, the French bacteriologist who first gave a clearly recognizable descrip- tion of this type of methane organism. Small spherical cells, occurring singly, in large, irregular masses, or in regular cysts of various sizes and forms. Non- motile. Stains readily with erythrosine. Gram-variable. Grows on calcium acetate enrichment media and ferments the acetate vigor- ously. Grows slowly on agar containing 2 per cent clear mud extract. Ferments acetic and butyric acids with production of methane in the presence of CO2. Ethyl and butyl alcohols not attacked. Does not utilize organic nitrogen. Obligate anaerobe. Grows best at 30° to 37°C. Sources: Garden soil, black mud con- taining HoS, feces of herbivorous ani- mals. Habitat : One of the most active meth- ane-producing organisms found in nature. * Appendixes T and II prepared by Experiment Station. Geneva, New York Prof. Robert S. Breed, New York State December, 1943. FAMILY MICROCOCCACEAE 249 Appendix II: The following genus is recognized by workers in the brewing industry. It includes species that present characters intermediate between Micrococcus, Sarcina and Sireptococctis . Many students prefer to regard these as species of Micrococcus (Hucker, N. Y. State Exper. Sta., Tech. Bui. 102, 1924, 5), of Sarcina (Mace, Traite pratique d. Bact., 4th ed., 1901, 460) or of Streptococcus (Shimwell, Sect. 670 in Hind, Brewing Science and Practice, New York, 1940). Others (Mees, Thesis, Delft, 1934) would include in the genus, the species described as Tetracoccus by Orla -Jensen (The Lactic Acid Bacteria, Copenhagen, 1919, 76). Genus B. Pediococcus Balcke. (Wchnschr. f. Brauerei, 1, 1884, 257.) Cocci occurring singh^, in pairs and tetrads. Xon-motile. No endospores. Gram- positive. Facultative anaerobes under favorable conditions, especially in acid media. Nitrites not produced from nitrates. Produce acidification and more or less clouding of wort and beer. Saprophytes. The tj'pe species is Pediococcus cerevisiae Balcke. 1. Pediococcus cerevisiae Balcke. (Ferment No. 7, Pasteur, Etudes sur la biere, Paris, 1876, 4; ySarcina Balcke, Wchnschr. f. Brauerei, /, 1884, 183; ibid., 1, 1884, 257 ; Merismopedia cerevisiae Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 348; Micrococcus cerevisiae Migula, Syst. d. Bakt., 2, 1900, 77; Sarcina cerevisiae Mace, Traits Pratique d. Bact., 4th ed.. 1901, 460.) From Latin, cerevisia, beer. Spheres: 1 to 1.3 microns, occurring singly, in pairs or tetrads. In acid media the latter prevail. Catalase negative. Non-motile. Gram-positive. No growth in alkaline media. Peptone, meat-e.xtract gelatin : White becoming yellowish to yellowish brown. No liquefaction. Wort gelatin with Ca-carbonate : White colonies, 2 to 3 mm; carbonate dissolved. Meat extract gelatin stab: Growth along stab, white raised surface growth. No liquefaction. Litmus milk: No growth. Potato: Scanty growth. Acid from glucose, fructose, maltose, sucrose. Wort and beer : Slight to moderately turbid growth, strong development on bottom of the flask. Hop sensitive, but may develop in heavily hopped beers under special conditions. Does not utilize urea. Nitrites not produced from nitrates. Facultative anaerobic. Killed at 60°C. in 8 minutes. Optimum temperature: 25°C. Source : Sarcina-sick beer. Habitat: Wort, beer and beer yeast. Additional species have been described from spoiled wort and beer which vary but slightly from the species first named and described by Balcke. These are listed below together with other species that have been placed in the genus. Pediococcus acidilactici Lindner. (Lindner, Wchnschr. f. Brauerei, 3, No. 23, 1887, see Cent. f. Bakt., 2, 1887, 342; also see Die Sarcina-Organismen der Gahrungsgewerbe, Lindner, Inaug. Diss., Berlin, 1888, 26, and Cent. f. Bakt., 4, 1888, 429; Micrococcus pseudocerevisiae Migula, Syst. d. Bakt., 2, 1900, 77; Mi- crococcus acidi-lactici Chester, Man. Determ. Bact., 1901, 88.) From spoiled mash. Pediococcus albus Lindner. (Die Sar- cina-Organismen der Gahrungsgewerbe, Lindner, Inaug. Diss., Berlin, 1887, 39; see Cent. f. Bakt., 4, 1888, ^29; Micrococ- cus pseudosarcina Migula, Sj^st. d. Bakt., 2, 1900, 92; Micrococcus albus Chester, Man. Determ. Bact., 1901, 97.) From spoiled beer. Pediococcus damnosus Claussen. 250 MANUAL OF DETERMINATIVE BACTERIOLOGY (Compt. rend. Trav. Labor, de Carlsberg, 6, 1906, 68; Streptococcus dajnnosus Shim- well and Kirkpatrick, Jour. Inst. Brew- ing, 45, 1939, 137.) From clear, spoiled beer. Pediococcus halophilus Mees. (Tetra- coccus No. 1, Orla-Jensen, The Lactic Acid Bacteria, 1919, 77; Mees, Thesis, Delft, 1934, 94.) From anchovy pickle. Pediococcus hennebergi Sollied. (Ztschr. Spiritusindus., 26, 1903, 491.) From spoiled beer. Pediococcus kochii Trevisan. (Mikro- kokkus in Wundsecreten bei Menschen, Koch; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 28.) Pediococcus pentosaceus Mees. (Tetra- coccus No. 2, Orla-Jensen, The Lactic Acid Bacteria, 1919, 78; Mees, Thesis, Delft, 1934, 94.) From yeast. Pediococcus perniciosus Claussen (loc. cit.). From clouded, spoiled beer. Pediococcus sarcinaeformis Rei chard. (Ztschr. f. d. ges. Brauwesen, 17, 1894, 257.) From spoiled beer. Pediococcus urinae equi Mees. (Pfer- deurinsarcina, von Huth, Alg. Ztg. f. Bierber. u. Malzfabr., 185, 968 and 981, 1885; ibid., 1886, 141; Mees, Thesis, Delft, 1934, 95.) From horse urine. Pediococcus violaceus (Klitzing) Trevi- san. {Merismopedia violacea Klitzing; Trevisan, I generi c le specie delle Bat- teriacee, Milan, 1889, 28.) Pedioplana haeckeli Wolff. (Cent. f. Bakt., II Abt., 18, 1907, 9.) Motile. From rotting beets. Placed in a new genus Pedioplana Wolff {loc. cit., 9). Streptococcus damnosus var. mucosus Shimwell. (Shimwell, Sect. 670, Hind, Brewing Science and Practice, New York, 1940.) From ropy beer. Appendix III*. The following species have been found in the literature and are listed here chiefly for their historical interest. Many are incompletely de- scribed, while many others are identical with previously described species. See Monographs by Winslow and Winslow, Sj'stematic Relationships of the Cocca- ceae, 1908 and Hucker, N. Y. State Exper. Sta., Tech. Buls. Nos. 99-103. Refer- ences are to Tech. Bui. 102. Ascococcus cantabridgensis Hankin. (Quoted from Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 165.) Mi- gula (Syst. d. Bakt., 2, 1900, 195) reports he is unable to find further reference to this organism and we likewise are unable to trace it. From the human mouth. Ascococcus gangrenosus Bevan. (Med. News, No. 1003, 1892, 375; Abst. in Cent, f. Bakt., 13, 1893, 796.) From a gan- grenous foot. Ascococcus vibrans van Tieghem. (Bui. Soc. Bot. France, 27, 1880, 150.) From water. Aurococcus tropicus Chalmers and O'Farrell. (1913, quoted from Castel- lani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 931.) Found in Ceylon in granulating ulcers of skin. Coccus carduus Heurlin. (Bakt. Un- ters. d. Keimgehaltes im Genitalkanale d. fiebernden Wochnerinnen. Helsing- fors, 1910, 136.) Anaerobic. From genital canal. Coccus caudatus Heurlin {loc. cit., 84). From genital canal. Coccus vaginalis Heurlin {loc. cit., 79). From genital canal. Galactococcus albus Guillebeau. (Land- wirtsch. Jahrb. d. Schweiz, 4, 1892, 27; Abst. in Cent. f. Bakt., 12, 1892, 101.) From milk from an inflamed udder. Galactococcxis fulvus Guillebeau {loc. cit.). From milk from an inflamed udder. Galactococcus versicolor Guillebeau {loc. cit.). From milk from an inflamed udder. Gyrococcus flaccidifex Glaser and Chap- man. (Science, 36, 1912, 219.) Isolated from the gypsy moth, Porthetria dispar. Jodococcus vaginatus Miller. (Miller, * Prepared for Prof. G. J. Hucker by Mrs. Eleanore Heist Clise, New York State Experiment Station, Geneva, New York, March, 1943. . FAMILY MICROCOCCACEAE 251 Mikroorganismen der Mundhohle, 1889, 54; Bacterium iogenum Baumgartner, Ergebnisse d. ges. Zahnheilk., Heft 2, 1910, 729; Abst. in Cent. f. Bakt., I Abt., Ref., 48, 1911, 621.) From the oral cavity. Merismopedia auranliaca Maggiora. (Giorn. Soc. Ital. d'Igiene, 11, 1889, 354; Abst. in Cent. f. Bakt., 8, 1890, 13.) From the normal skin of the human foot. Micrococcu!^ achrous Migula. (No. 16, Lembke, Arch. f. Hyg., 26, 1896, 310; Migula, Syst. d. Bakt., 2, 1900, 201.) From feces. Winslow and Winslow (Sys- tematic Relationships of the Coccaceae, 1908, 224) state that this species is appar- ently a synonym of Micrococcus candi- cans Fliigge. Micrococcus acidi lactici Marpmann. (Erganzungsheft d. Cent, f . allg. Gesund- heitspflege, 2, 1886, 22.) Found in fresh milk. Micrococcus acidovorax MuUer-Thur- gau and Osterwalder. (Cent. f. Bakt., II Abt., 86, 1913, 236.) From wine. Hucker {loc. cit., 6) considers this a synonym of Micrococcus luteus Cohn or Micrococcus varians Migula. I\[icrococcus acne Hollaml. (Jour. Bact., S, 1920, 223; Staphylococcus acne Holland, ibid., 225; see Micrococcus cittis communis Sabouraud.) Micrococcus (Staphylococcus) acridi- cida Kufferath. (Ann. de CJembloux, 27, 1921, 253.) Isolated from diseased locusts from Greece. Resembles Micro- coccus aureus Zopf. Micrococcus aerius Chester. (Xo. 49, Conn, Storrs Agr. Exp. Sta. 7th Ann. Rept., 1895, 81; Chester, Man. Determ. Bact., 1901, 104.) From dust. Hucker {loc. cit., 12) states that this species appears to be identical with Micrococcus aureus Zopf. Micrococcus aerogenes Miller. (Miller, Deutsche med. Wchnschr., 12, 1886, 119; not Micrococcus aerogenes Bergey et al., Manual, 1st ed., 1923, 70.) P>om the alimentary canal. Micrococcus agilis albus Catterina. (Cent. f. Bakt., I Abt., Orig., 34, 1903, 108.) Found in septicemia of rabbits. Motile with one or two flagella. Micrococcus alhatus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 479.) From the intestine of a woodpecker {Picus major). Winslow and Winslow (Systematic Relationships of the Coc- caceae, 1908, 199) state that this species appears to be a synonym of M icrococcus albus Schroeter; while Hucker (N. Y. Agr. Exper. Sta., Tech. Bull. 102, 19) regards it as a synonym of Micrococcus freudenreichii Guillebeau or M icrococcus ureae Cohn. Micrococcus albescens Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 76.) From cheese. Winslow and Wins- low (loc. cit., 199) state that this species appears to be a synonym of Micrococcus albus Schroeter; while Hucker (loc. cit., 19) regards it as a synonym of Micrococ- cus freudenreichii Guillebeau or of Mi- crococcus ureae Cohn. Micrococcus albidus Losski. (Losski, Inaug. Diss., Dorpat, 1893, 55; not Micrococcus albidus Henrici, see Micro- coccus subniveus below; not Micrococcus albidus Roze, Compt. rend. Acad. Sci. Paris, 122, 1896, 750.) From soil. Hucker (loc. cit., 19) regards this species as a synonym of Micrococcus freuden- reichii Guillebeau or Micrococcus ttreae Cohn. Micrococcus albocereus Migula. (Sta- phylococcus cereus albus Passet, Unter- such. ii. d. Aetiol. d. eiterigen Phlegmone d. Menschen, Berlin, 1885, 53, and Fort- schr. d. Med., 3, 1885; Micrococcus cereus albus Fltigge, Die Mikroorganismen, 2 Aufl., 1886, 182; Staphylococcus cereus Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 32; Migula, Syst. d. Bakt., 2, 1900, 56; Staphylococcus cereus-albus Holland, Jour. Bact., 5, 1920, 225.) From human pus, also from water. Winslow and Winslow (Systematic Rela- tionships of the Coccaceae, 1908, 205) consider this a synonym of M icrococcus 252 MANUAL OF DETERMINATIVE BACTERIOLOGY candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcxis albus Frankland and Frankland. (Phil. Trans. Roy. Soc, Lon- don, 178, B, 1888, 264; not Micrococcus albus Matzuschita, Cent. f. Bakt., I Abt., 29, 1901, 382; not Micrococcus albus Buchanan, Veterinary Bacteri- ology, 1911, 196; not Micrococcus albus Mace, Traits Pratique de Bact., 6th ^d., 1912, 605.) From air. Resembles Micrococcus candicans. Micrococcus albus II Maggiora. (Cent. f. Bakt., 8, 1890, 13.) See Micro- coccus opalescens De Toni and Trevisan. From the skin of the human foot. Micrococcus ampins Migula. (Grau- weisser Diplococcus, Bumm, Der Mikro- org. d. gonorrh. Schleimhauterkrank., 1 Aufl., 1885, 17; Micrococcus albicans ampins Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 183; Neisseria albicans Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 32; Diplococcus albicans amplus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 24; Migula, Syst. d. Bakt., 2, 1900, 118 ; Micrococcus albicans Chester, Man. Determ. Bact., 1901, 80.) From vaginal secretions. Hucker (loc. cit., 15) considers this species identical with Micrococcus albus Schroeter. Micrococcus ampullaceus Kern. (Arb. bakt. Inst. Karlsruhe, ^ , Heft 4, 1897, 477.) From the intestine of a dove (Columba oenas). Hucker (loc. cit., 19) considers this a synonym of Micrococcus freuden- reichii Guillebeau or Micrococcus ureae Cohn. Micrococcus annulatus Kern {loc. cit., 490). From the stomach contents of the hedge sparrow (Passer montanus) and the intestine of the rock dove {Columba livia). Winslow and Winslow (Syste- matic Relationships of the Coccaceae, 1908, 216) consider this species a synonym of Micrococcus flavus Lehmann and Neu- mann. Micrococcus aquatilis Bolton. (Zschr. f. Hyg., /, 1886, 94; not Micrococcus aquatilis Chester, see below.) From water. Winslow and Winslow {loc. cit., 224) state that this species is apparently a synonym of Micrococcus candicans Flugge. Micrococcus aquatilis Chester. {Mi- crococcus aquatilis invisibilis Vaughan, Amer. Jour. Med. Sci., /04, 1892, 183; Chester, Man. Determ. Bact., 1901, 88.) From water. Winslow and Winslow {loc. cit., 224) state that this species is ap- parently a synonym of Micrococcus candi- cans Flugge. Micrococcus aquatilis albus. (Quoted from Toporoff, Protok. d. Kaiserl. kau- kasisch Mediz. Gesellsch., 1892, No. 21; Abst. in Cent. f. Bakt., 13, 1893, 487.) From water. Micrococcus aquatilis flavus. (Quoted from Toporoff, loc. cit.) From water. Micrococcus aqueus Migula. (No. 25, Lembke, Arch. f. Hyg., 26, 1896, 317; Migula, Syst. d. Bakt., 2, 1900, 204.) From feces. Winslow and Winslow {loc. cit., 184) state that this species is appar- ently a synonym of Micrococcus aureus Zopf, while Hucker {loc. cit., 15) regards this as a synonym of Micrococcus albus Schroeter. Micrococcus arbor escens lactis Conn. (Conn, Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 46; Micrococciis lactis ar- borescens Conn, Esten and Stocking, Storrs Agr. Exp. Sta. 18th Ann. Rept., 1907, 110.) From milk. Hucker {loc. cit., 21) regards this as a synonym of Micrococcus candidus Cohn or Micrococ- cus epidermidis Hucker. Micrococciis argenteus Migula. (No. 27, Lembke, Arch. f. Hyg., 26, 1896, 317; Migula, Syst. d. Bakt., 2, 1900, 206.) From feces. Winslow and Winslow {loc. cit., 184) state that this species is appar- ently a synonym of Micrococcus aureus Zopf, while Hucker {loc. cit., 10) con- siders it a synonj'm of Micrococcus con- glomcratus Migula. Micrococcus ascoformans J ohne . {Zoo- gloea pulmonis equi Bollinger, Arch. f. path. Anat., 49, 1870, 583; Discomyces equi Rivolta, Giorn. di Anat. e Fisiolog., FAMILY MICROCOCCACEAE 253 10, 1884; Johne, Ber. u. d. Veterin. im Konigr. Sachsen, Jahr 1885, 47; Ascococ- cus johnei Cohn in letter to Johne, Deutsche Ztschr. f. Thiermed., 12, 1886, 210; Micrococcus botryogenus Rabe, Deut. Ztschr. f. Thiermed., 12, 1886, 137; Botryomyces equi Bollinger, Deut. Ztschr. f. Thiermed., 13, 1887, 176; Botryococcus ascoformans Kitt, Cent. f. Bakt., 3, 1888, 247; BoUingera equi Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 26; Staphylococcus ascoformans Ford, Textb. of Bact., 1927, 424; Staphylococcus aureus var. equi Hauduroy et al.. Diet. d. Bact. Path., 1937, 504.) Causes botryomycosis in horses. Lehmann and Neumann (Bakt. Diag., 7 Aufl., 2, 1927, 291) consider this a form of Micrococcus aureus Zopf ; while Hucker {loc. cit., 15) regards this as a form of Micrococcus albus Schroeter. Micrococcus ascoformis Fermi. (Arch, f. Hyg., 10, 1890, 10.) Presumably in- tended for Micrococcus ascoformans Johne. Micrococcus asper Migula. (Seibert, Inaug. Diss., Wtirzburg, 1894, 12; Migula, Syst. d. Bakt., 2, 1900, 82.) From a hairbrush. Winslow and Winslow (loc. cit., 205) consider this species to be a synonym of Micrococcus candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcus aurantiacus-sorghi Bruyn- ing. (Arch. Neer. Sci. Exact, et Nat., /, 1898, 297; Streptococcus aurantiacus Chester, Man. Determ. Bact., 1901, 69.) From sorghum. Micrococcus aureus Chester. (Micro- coccus cremoides aureus Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 349; Chester, Manual Determ. Bact., 1901, 99.) From dust. Regarded by Dyar as a varietal form of Micrococcus cremoides Zimmermann. Winslow and Winslow (loc. cit., 184) consider this species a synonym of Mi- crococcus aureus Zopf. Micrococcus aureus lactis Conn. (Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 36.) From milk. This seems to be identical with Micrococcus lactis aureus A, Conn, Esten and Stocking, Storrs Agr. Exper. Sta. 18th Ann. Rept., 1907, 119. Hucker (loc. cit., 9) regards this species as identical in part with Micrococ- cus jlavus Lehmann and Neumann and with Micrococcus conglotneratus Migula. Micrococcus (Sarcina) baccatus Migula. (No. 18, Lembke, Arch. f. Hyg., ^6, 1896, 311; Migula, Syst. d. Bakt., 2, 1900, 202.) From feces. Winslow and Winslow (loc. cit., 232) state that this is a yellow, gela- tin-liquefying sarcina, apparently a syn- onym of Sarcina flava De Bary . Hucker (loc. cit., 10) considers this a synonym of Micrococcus conglomeratus Migula. Micrococcus badius Lehmann and Neu- mann. (Bakt. Diag., 1 Aufl., 2, 1896, 163.) Received from the Krai collection as Sarcina lutea. Winslow and Winslow (loc. cit., 216) consider this a synonym of Micrococcus flavus Lehmann and Neu- mann. Micrococcus baregensis purpureus Ro- bine and Hauduroy. (Compt. rend. Soc. Biol., Paris, 98, 1928, 25.) From hot sulfur springs at Bareges. Micrococcus beigelii (Rabenhorst) Mi- gula. (Pleurococcus beigelii Kiichen- meister and Rabenhorst, Hedwigia, 1867, No. 4; Sclerotium beigelianum Hallier, 1868; Zoogloea beigeliana Eberth, 1873; Hyalococcus beigelii Schroeter, Kryptog.- Flora V. Schlesien, 3, 1, 1886, 152; Chlamydatomus beigelii Trevisan, Rendi- conti Reale Inst. Lombardo de Sci. e Lett., Ser. II, 12, 1879, 22; Migula, Syst. d. Bakt., 2, 1900, 193; Trichosporum beigelii Vuillemin, 1901.) From human hair. Micrococcus beri-beri Pekelharing. (Pekelharing, Weekblad v. h. Ned. Tijdschr. v. Geneesk., No. 25; also Pekelharing and Winkler, Deut. med. Wchnschr., No. 39, 1887, 845; Neisseria loinkleri Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 32.) Con- sidered the cause of beri-beri by Pekel- haring. Winslow and Winslow (loc. cit., 184) state that this is apparently a synonym of Micrococcus aureus Zopf; while Hucker (loc. cit., 11) considers 254 MANUAL OF DETERMINATIVE BACTERIOLOGY this a synonym of Micrococcus cilreus Migula. Micrococcus hicolor Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 485.) From the intestine of a dove {Columba oenas). Hucker (loc. cit., 21) considers this a synonym of Micrococcus candidus Cohn or of Micrococcus epidermidis Hucker. Micrococcus billrothii (Cohn) Migula. (Ascococcus billrothii Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 3, 1875, 151; Migula, Syst. d. Bakt., 2, 1900, 195.) Found in putrefying meat infusion. Micrococcus biskra Heydenreich. (Heydenreich, Ausgabe d. Haupt Med.- Verhalt., St. Petersburg, 1888; see Cent, f. Bakt., 5, 1889, 163; Staphylococcus biskrae Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 32; Micrococ- cus heydenreichii Chester, Man. Determ. Bact., 1901, 91.) Found in ulcers in an Oriental skin disease. Winslow and Winslow (loc. cit., 184) state that this is apparently a synonym of Micrococcus aureus Zopf; while Hucker (loc. cit., 11) considers it a synonym of Micrococ- cus cilreus Migula. Micrococcus boleti Passerini. (Erbar. crittogam. Italiano, II ser.. No. 1199; quoted from Trevisan, I generi c le specie delle Batteriacee, Milan, 1889, 34.) Saprophytic on a fungus (Boletus edulis). Micrococcus bombycis (Naegeli) Cohn. (Panhistophyton ovatum Lebert, Jahres- ber. i'l. d. Wirksamkeit d. Vereins z. Beford. d. Seidenbaues f. Brandenburg im Jahre 1856-57, 28; and Berliner En- tomolog. Ztschr., 1858; Nosema bombycis Naegeli, Botan. Sect. d. 33 Versammlg. d. Naturf. u. Aerzte in Bonn, 1857, 160; and Botan. Zeitg., 1857, 760; Microzyma bombycis Bechamp, Compt. rend. Acad. Sci., Paris, 6^, 1867, 1045; Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 1872, 165; Micrococcus ovatus Winter, in Raben- horst, Krypt. Flora v. Deutschl., Oes- terr. u. d. Schweiz, 2 Aufl., 1, 1884, 47; Streptococcus bombycis Zopf, Die Spalt- pilze, 2 Aufl., 1884, 52.) Found in the blood and organs of diseased silkworms (Bombyx mori). Micrococcus boreus Issatchenko. (Re- cherches sur les Microbes de I'Ocedn Glacial Arctique, Petrograd, 1914, 144.) From sea water. Micrococcus bovinus Migula. (Micro- coccus der Lungenseuche der Rinder, Poels and Nolen, Fortschr. d. Med., 1886, 217; Migula, Syst. d. Bakt., 2, 1900, 67.) From the lungs of diseased cattle. Hucker {loc. cit., 22) regards this a synonym of Micrococcus candidus Cohn or of Micrococcus epidermidis Hucker. Micrococcus bovis Migula. (Hemato- coccus, Babes, Compt. rend. Acad. Sci., Paris, 107, 1888, 692 and 110, 1890, 800 and 975; also see Arch. f. path. Anat., 115, 1889; Neisseria babesi Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 32; Haematococcus bovis Eisenberg, Bakt. Diag., 3 Aufl., 1891, 271 ; Migula, Syst. d. Bakt., 2, 1900, 85.) From the blood and organs of cattle. Micrococcus burchardti Trevisan. (Coccus bei keratitis phlyctaenulosa, Burchardt, Cent. f. Bakt., 1, 1887, 392; Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 33.) Pathogenic. From the cornea of a rabbit. Micrococcus butyri (v. Klecki) Migula. (Diplococcus butyri von Klecki, Cent, f. Bakt., 16, 1894, 358; Migula, Syst. d. Bakt ., ^, 1900, 216. ) From rancid butter. Winslow and Winslow (loc. cit., 220) con- sider this a synonym of Micrococcus luteus Cohn. Micrococcus butyrlfluorescens Teichert. (Inaug. Diss., Jena, 1904; Abst. in Cent, f. Bakt., II Abt., IS, 1904, 561.) From milk. Exhibits a green fluorescence. Micrococcus butyricus (von Klecki) Migula. (Tetracoccus butyricus von Klecki, Cent. f. Bakt., 15, 1894, 360; Migula, Syst. d. Bakt., 2, 1900, 216.) From rancid butter. Winslow and Win- slow (loc. cit., 220) consider this a syn- onym of Micrococcus luteus Cohn. FAMILY MICROCOCCACEAE 255 Micrococcus calco-aceticus Beijerinck. (Proc. Sect. Sci., Kon. Akad. v. Weten- schappen, 13, 1911, 1066; Abst. in Cent, f. Bakt., II Abt., 31, 1912, 290.) Occurs commonly in soils. Micrococcus campeneus Orme. (Jour. Trop. Med. and Hyg., 11, 1908, No. 10, May 15; Abst. in Cent. f. Bakt., I Abt., Ref., 43, 1909, 299.) Micrococcus candicans Fltigge. (Die Mikroorganismen, 2 Aufl., 1886, 173; Albococcus candicans Winslow and Rogers, Jour. Inf. Dis., 3, 1906, 544; Staphylococcus candicans Holland, Jour. Bact., S, 1920, 225.) From air, water and milk. Hucker (loc. cit., 22) regards this a synonym of Micrococcus candidus Cohn or of Micrococcus epidermidis Hucker. For a description of this spe- cies, see Bergey et al., Manual, 5th ed., 1939, 255. Micrococcus canescens Migula. (Mi- crococcus No. 4, Adametz, Landwirtsch. Jahrb., 18, 1889, 240; Migula, Syst. d. Bakt., 2, 1900, 51; Albococcus cariescens Winslow and Rogers, Jour. Inf. Dis., 8, 1906, 544; Staphylococcus canescens Hol- land, Jour. Bact., 5, 1920, 225.) From Emmenthal cheese. Winslow and Win- slow {loc. cit., 224) state that this is apparently a synonym of Micrococcus candicans Fliigge. Micrococcus capillorum (Buhl) Tre- visan. (Zoogloea capillorum Buhl, Ztschr. f. ration. Med., II Reihe, U, 18-, 356; Palmella capillorum Ktihn, Abhandl. d. Naturf. Ges. zu Halle, 9, Heft 1, 18-, 62; Palmellina capillorum Rabenhorst, Flor. Eur. Alg., 3, 1856 (?), 35;Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33.) From the skin. Con- sidered pathogenic. Micrococcus capsaformans Jamieson and Edington. (Brit. Med. Jour., 1, June 11, 1887, 1262; Micrococcus capriformis (sic), Abst. in Cent. f. Bakt., 2, 1887, 223.) From the scales and blood of scarlet fever patients. Not pathogenic. Micrococcus carbo Renault. (Compt. rend. Acad. Sci., Paris, 123, 1896, 935.) Micrococcus carneus Zimmermann. (Roter Coccus, Maschek. Bakt. Unter- such d. Leitmeritz. Trinkwassers, No. 5, 1887, 60; Zimmermann, Die Bakt. unserer Trink- u. Nutzwasser, Chem- nitz, I Reihe, 1890, 78.) From water. Hucker {loc. cit., 25 and 26) regards this species as identical with Micrococcus roseus Fliigge or Micrococcus cinna- bar eus Fliigge. Micrococcus' carnicolor Frankland and Frankland. (Phil. Trans. Roy. Soc. London, 178, B, 1888, 263; not Micrococ- cus carnicolor Kern, see Micrococcus subcarneus below.) From air. Hucker (loc. cit., 25) states that this species may be identical with Micrococcus roseus Fliigge. Micrococcus carniphilus Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 10.) From a meat extract. Micrococcus casei amari edamicus Raamot. (Inaug.Diss.,K6nigsberg, 1906; Abst. in Cent. f. Bakt., II Abt., 18, 1907, 348.) From pasteurized skim milk. Micrococcus castellanii Chalmers and O'Farrell. (Ann. Trop. Med. and Para- si tol., 7, 1913, 528; Rhodococcus castellanii Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 2102.) Found in the red variety of trichomycosis axillaris, a tropical disease. Micrococcus cartharinensis Issat- chenko. (Recherches sur les Microbes de I'Oce^n Glacial Artique, Petrograd, 1914, 148.) From sea water. Micrococcus cellaris (Schroeter) Mi- gula. {Leucocystis cellaris Schroeter, Kryptog. Flora v. Schlesien, 3, 1886, 152; Migula, Syst. d. Bakt., 2, 1900, 195.) From a coating on the walls of damp cellars and mines. Micrococcus centropunctatus Issat- chenko. (Recherches sur les Microbes de rOcedn Glacial Arctique, Petrograd, 1914, 146.) From sea water. Micrococcus cerasinus ^ligula. (]Mi- crococcus aus roter Milch, Keferstein; Cent. f. Bakt., I Abt., 21, 1897, 177; .1/z- crococcus cerasinus lactis Helm, Lehrb. 256 MANUAL OF DETERMINATIVE BACTERIOLOGY d. Bakt., 2 Aufl., 1898, 299; Migula, Syst. d. Bakt., £, 1900, 170; not Micro- coccus cerasinus Lehmann and Neumann, Bakt. Diag., 1 AuH., 2, 1896, 179; Micro- coccus kefersteinii Chester, Man. De- term. Bact., 1901, 107.) From red milk. Hucker (loc. cit., 26) regards this species as identical with Micrococcus cinna- bareus Fliigge. Micrococcus cereus Migula. (Staphy- lococcus cereus flavus Passat, Unter- suchungen liber die Aetiologie der eiteri- gen Phlegmone des Menschen, 1885, 53; Micrococcus cereus flavus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 182; Staphylococcus passeti Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 32; Migula, Syst. d. Bakt., 2, 1900, 126; Staphylococcus cereus-flavus Hol- land, Jour. Bact., 5, 1920, 225.) From pus. Winslovv and Winslow {loc. cit., 220) consider this species identical with Micrococcus luteus Migula. For a de- scription of this organism, see Bergey et al., Manual, 5th ed., 1939, 241. Micrococcus cereus aureus Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 347.) Obtained as Staphylococcus cereus aureus from Krai's laboratory; also found in air. Micrococcus cerinus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 84.) From Swiss cheese. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus chersonesia Corbet. (Quart. Jour. Rubber Research Inst., Malaya, 2, 1930, 150.) From the latex of the rubber tree {Hevea brasiliensis) . For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 258. Micrococcus chinicus Enimerling and Abderhalden. (Cent. f. Bakt., II Abt., 10, 1902, 337.) Putrefying meat. Micrococcus chlorinus Cohn. (Griin- gelber Coccus, Maschek, Jahresber. d. Kom.- Oberrealschule zu Leitmeritz, 1887, 66; Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 1872, 155.) From water and dust. Hucker {loc. cit., 10) considers this a synonym of Micrococcus conglom- eratus Migula. Micrococcus chromidrogenus citreus Trommsdorff. (Miinch. Med. Wochn- schr., 1904, No. 29, 1286; Abst. in Cent, f. Bakt., I Abt., Ref., 37, 1905, 60.) Isolated from a case of chromidrosis of the axilla. Micrococcus chromidrogenus ruber Trommsdorff {loc. cit.). Isolated from a case of chromidrosis of the axilla. Micrococcus chromoflavus Huss. (Cent. f. Bakt., II Abt., 19, 1907, 520.) From cheese Micrococcus chryseus Frankland and Frankland. (Phil. Trans. Roy. Soc. London, 178, B, 1888, 268.) From dust. Winslow and Winslow {loc. cit., 184) state that this species is apparently a synonym of Micrococcus aureus Zopf. Micrococcus cinnabarinus Zimmer- mann. (Die Bakt. unserer Trink- u. Nutzwasser, Chemnitz, I Reihe, 1890, 76.) From water. Hucker (Zoc. czi., 26) regards this species as identical with Micrococcus cinnabareus Fliigge. Micrococcus cirrhiformis Migula. (Ranken Coccus, Maschek, Jahresber. (1. Kom.- Oberrealschule in Leitmeritz, 1887, 66; Migula, Syst. d. Bakt., 2, 1900, 53.) From water. Hucker {loc. cit., 22) considers this a synonym of Micrococcus candidus Cohn or of Micrococcus epider- midis Hucker. Micrococcus citreus I and //, Maggiora. (Giorn. Soc. Ital. d'Igiene, 11, 1889, 354; Abst. in Cent. f. Bakt., 8, 1890, 13; not Micrococcus citreus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 36; not Micrococcus citreus Migula, Syst. d. Bakt., 2, 1900, 147; not Micrococcus citreus Winslow and Winslow, Systematic Relationships of the Coccaceae, 1908, 218.) From the normal skin of the foot. Micrococcus citreus granulatus Freund. (Inaug. Diss., Erlangen, 1893, 27; Abst. in Cent. f. Bakt., 16, 1894, 641 ; Micrococ- cus granulatus Bazarewski, Cent. f. Bakt., II Abt., 15, 1905, 7; not Micrococ- cus granulatus Weiss, Arb. bakt. Inst. FAMILY MICROCOCCACEAE 257 Karlsruhe, 2, Heft 3, 1902, 197.) From the oral cavity. Hucker Q.oc. cit., 9) regards this as a synonym of Micrococcus flavus Trevisan. Micrococcus citreus lactis Conn. (Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900,. 40.) From milk. Hucker {loc. cit., 10) considers this a synonym of Micrococcus conglomeratus ]Migula. Micrococcus citreus rigensis Bazarew- ski. (Cent. f. Bakt., II Abt., 15, 1905, 5.) From dust. Micrococcus citrinus Migula. (Diplo- coccus citreus liquefaciens Unna and Tommasoli, Monats. f. prakt. Dermatol - ogie, 9, 1889, 56; Migula, Syst. d. Bakt., 2, 1900, 150; Micrococcus tommasoli Ches- ter, Man. Determ. Bact., 1901, 101; Mi- crococcus citreus liquefaciens Winslow and Winslow, Systematic Relationships of the Coccaceae, 1908, 216.) From hu- man skin in a case of eczema. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevi- san. Micrococcus claviformis von Besser. (Beitr. z. path. Anat. u. z. allgem. Path., 6, 1889, 340; see Cent. f. Bakt., 7, 1890, 152.) Found once in nasal secretions. Micrococcus coccineus Migula. (Mi- crococcus No. VI, Adametz,Landwirtsch. Jahrb., 18, 1889, 242; Migula, Syst. d. Bakt., 2, 1900, 174.) From Emmenthal cheese. Hucker (loc. cit., 26) regards this species identical with Micrococcus cinnabareus Fltigge. Micrococcus coli brevis Lehmann. (Lehmann, Inaug. Diss., Miinchen, 1903; Abst. in Cent. f. Bakt., I Abt., Ref., 36, 1905, 688.) From feces of in- fants. Micrococcus communis lactis Conn. (Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 48.) From milk. Hucker (loc. cit., 19) considers this a synonym of Micrococcus freudenreichii Guillebeauor of Micrococcus ureae Cohn. Micrococcus commensalis (Turro) Mi- gula. (Diplococcus commensalis Turro, Cent. f. Bakt., 16, 1894, 1; Migula, Syst. d. Bakt., 2, 1900, 125.) From sputum. Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus commutatus De Toni and Trevisan. {Micrococcus albus I or Mi- crococcus albus fluidificans Maggiora, Giorn. Soc. Ital. d'Igiene, 11, 1889, 350; De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1079.) Micrococcus concentricus Zimmermann. (Die Bakt. unserer Trink- u. Nutz- wjisser, Chemnitz, I Reihe, 1890, 86.) From water. Winslow and Winslow (loc. cit., 224) state that this is appar- ently a synonym of Micrococcus candicans Fltigge. Micrococcus confluens Kern. (Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1897, 494.) From the stomach and intestine of the starling {Sturnus vulgaris) and the finch {Fringella carduelis). Win- slow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus conjunctivae Migula. {Micrococcus liquefaciens conjunctivae Gombert, Recherches experimentales sur les microbes des conjunctives, Mont- pellier and Paris, 1889; Migula, Syst. d. Bakt., 2, 1900, 115.) From normal hu- man conjunctiva. Hucker {loc. cit., 15) considers this a synonym of Micrococcus albus Schroeter. Micrococcus conjunctividis Migula. {Micrococcus flavus conjunctivae Gom- bert, Recherches experimentales sur les microbes des conjunctives, Montpellier and Paris, 1889; Migula, Syst. d. Bakt., 2, 1900, 141.) From normal human con- junctiva. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan; while Hucker {loc. cit., 11) regards it as a synonym of Micrococcus citreus Migula. Micrococcus conoideus Migula. (Staph- ylococcus salivarius pyogenes Biondi, Ztschr. f. Hyg., 2, 1887, 227 ; Staphylococ- cus sialopyus Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 258 MANUAL OF DETERMINATIVE BACTERIOLOGY 32; Micrococcus salivarius -pyogenes Freire, Memoire sur la bacteriologie, pathog^nie, traitement et prophylaxie de la fievre jaune. Rio de Janiero, 1898; Abst. in Cent. f. Bakt., I Abt., 26, 1899, 741; Staphylococcus pyogenes salivarius, quoted from Goadby, Trans, of Odon- tolog. Society, June, 1899, see Abst. in Cent. f. Bakt., I Abt., Ref., 31, 1902, 493; Migula, Syst. d. Bakt., 2, 1900, 102.) From saliva. Hucker {loc. cit., 12 and 15) regards this as a synonym of Micro- coccus aureus Zopf or of Micrococcus albus Schroeter. Micrococcus corallinus Cantani. (Can- tani. Cent, f . Bakt., I Abt., 23, 1898, 311 ; Rhodococcus coralinus (sic) Levine and Soppeland, Iowa State Coll. Engineer- ing Exp. Sta. Bui. 77, 1926, 22.) From dust. Hucker {loc. cit., 25) considers this a synonym of Micrococcus roseus Fliigge. Levine and Soppeland (loc. cit.) regard this as a synonym of Rhodococcus fulvus Winslow and Rogers. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 253. Micrococcus coralloides Zimmermann. (Die Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 72.) From water. Winslow and Winslow (loc. cit., 199) state that this species appears to be a synonym of Micrococcus albus Zopf; while Hucker (loc. cit., 17) considers it a synonym of Micrococcus caseolyticus Evans. Micrococcus corrugatus Migula. (Merismopedia mesentericus corrugatus Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 355; Migula, Syst. d. Bakt., 2, 1900, 161.) From dust. Winslow and Winslow (loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus coryzae (Hajek) Migula. (Diplococcus coryzae Hajek, Berliner klin. Wochnschr., No. 33, 1888; Migula, Syst. d. Bakt., 2, 1900, 63.) From secre- tions in acute catarrh. Winslow and Winslow (loc. cit., 205) consider this a synonym of Micrococcus candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcus cremoides Zimmermann. (Die Bakt. unserer Trink- u. Nutzwasser, Chemnitz, I Reihe, 1890, 74.) From water. Winslow and Winslow (loc. cit., 216) consider this a synonym of Micro- coccus flavus Trevisan; while Hucker (loc. cit., 10) considers it a synonym of Micrococcus conglomeratus Migula. Micrococcus cremoides albus Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 350). From dust. Regarded by Dyar as a white form of Micrococcus cremoides Zimmermann. Micrococcus cremorisviscosi (Hammer and Cordes) Bergey et al. (Staphylococcus cremoris-viscosi Hammer and Cordes, Jour. Dairy Sci., 3, 1920, 291; Bergey et al.. Manual, 3rd ed., 1930, 86.) From ropy milk. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 254. Micrococcus crepuscidum (Ehrenberg) Cohn. (Monas crepuscidum Ehrenberg, Abhandl. d. Berliner Akad., 1830, 74 and 1832, 57 ; Cohn, Beitr. z. Biol. d. Pflanzen, 1 , Heft 2, 1872, 160.) De Toni and Trevi- san (in Saccardo, Sylloge Fungorum, 8, 1889, 1082) list the following as synonyms of this species; Protococcus nebidosus Kiitzing, Linneae, 8, 1833, 365; Crypto- coccus nebulosus Kiitzing, Phycol. gener., 1845, 147; Cryptococcus natans Kiitzing, Spec. Alg., 1849, 146. Micrococcus cretaceus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 65.) From cheese. Winslow and Wins- low (loc. cit., 224) state that this is appar- ently a synonym of Micrococcus candicans Fliigge. Micrococcus cristatus Glage. (Ztschr. f. Fleisch- u. Milch-hygiene, 10, 1900, 145.) From the surface of wurst and similar meat products. Micrococcus cruciformis Freire. (Compt. rend. Acad. Sci., Paris, 128, 1899, 1047.) From the stamens and pistils of Hibiscus. Micrococcus cumidatus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 497; not Micrococcus cuinulatus Chester, FAMILY MICROCOCCACEAE 259 see Micrococcus tenuissimus INIigula.) From the stomach and intestine of the yellow-hammer {Emberiza citrinella) and of the finch {Fringella carduelis). Hucker {loc. cit., 25) regards this as a synonym of Micrococcus roseus Fliigge. Micrococcus cupularis Migula. (No. 29, Lembke, Arch. f. Hyg., 29, 1897, 331 ; Migula, Syst. d. Bakt., 2, 1900, 211.)From feces. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococ- cus flavus Trevisan. Micrococcus cupuliformis Migula. (No. 19, Lembke, Arch. f. Hyg., 29, 1897, 325; Migula, Syst. d. Bakt., 2, 1900, 213.) From feces. Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus curtissi Chorine. (Chor- ine, Internat. Corn Borer Invest. Chi- cago, 2, 1929, 48.) From diseased larvae of the corn borer (Pyrausta nubilalis). Also virulent to larvae of the flour moth {Ephestia kuhniella) and of the bee moth (Galleria mellonella). Micrococcus cutis communis Sabouraud. (Sabouraud, Ann. d. derma tol. et syphil., 1896, Heft 3; Abst. in Cent. f. Bakt., I Abt., 20, 1896, 249; Staphylococcus cutis communis Sabouraud, Practique Derma- tologique, /, 1903, 714.) From human skin especially in alopecia areata, certain types of eczema and acne. May be the same as Micrococcus epidermidis Hucker. Micrococcus cyaneus (Schroeter) Cohn. (Bacteridium cyaneum Schroeter, Beitr. z. Biol. d. Pflanzen, /, Heft 2, 1872, 122 and 126; Cohn, ibid., 156; Nigrococcus cyaneus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 932.) From dust and water. Micrococcus cyanogenus Pammel and Combs. (Proc. Iowa Acad. Sci., 3, 1895, 136; see Abst. in Cent. f. Bakt., II Abt., 2, 1896, 764.) From milk. Micrococcus cyclops Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 69.) From Swiss cheese. Winslow and Winslow (loc. cit., 224) state that this is appai'ently a synonym of Micrococcus candicans Fliigge. Micrococcus cystiopoeus Miiller-Thur- gau. (Cent. f. Bakt., II Abt., 20, 1908, 464.) From wine. Micrococcus cytophagus Merker. (Cent. f. Bakt., II Abt., 31, 1912, 589.) Found on the leaves of Elodea. Utilizes cellulose. Stanier (Bact. Rev., 6, 1942, 150) thinks that these micrococci were microcysts of Sporocytophaga spp. Micrococcus dantecH Chester. (Coc- cus du rouge de morue, Le Dantec, Ann. Past. Inst., 5, 1891, 662; Chester, Man. Determ. Bact., 1901, 106.) From red salted codfish. Hucker (loc. cit., 25) considers this a synonym of Micrococcus roseus Fliigge. Micrococcus decalvens (Thin) Schroe- ter. (Bacterium decalvens Thin, Monats. f. prakt. Dermatol., No. 28, 1885; Schroe- ter in Cohn, Kryptog. -Flora v. Schlesien, 3, 1, 1886, 149.) From hair follicles in alopecia areata. Micrococcus decipiens Trevisan. (Bac- terie de Pair, Cornil and Babes, Les Bacteries, 1885, 124; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 34.) From dust. Micrococcus (Streptococcus ?) decolor Migula. (No. 22, Lembke, Arch. f. Hyg., 26, 1896, 314; Migula, Syst. d. Bakt., 2, 1900, 203.) From feces. Hucker (loc. cit., 17) considers this a synonym of Micrococcus caseolyticus Evans. Micrococcus deformans Crowe. (Brit. Med. Jour., Nov. 27, 1920, 815; Abst. in Cent. f. Bakt., I Abt., Ref., 73, 1922, 84.) From cases of arthritis. A form of Micrococcus pyogenes albus according to Lehmann and Neumann (Bakt. Diag., 7 Aufl., 2, 1927, 293). Micrococcus delacourianus Roze. (Compt. rend. Acad. Sci., Paris, 123, 1896, 613 and 1323.) From dry rotting potatoes. Micrococcus dendroporthos Ludwig. (Cent. f. Bakt., 10, 1891, 10.) From the bark of poplar trees (Poprdus sp.). 260 MANUAL OF DETERMINATIVE BACTERIOLOGY Micrococcus denitrificans Beijerinck. (Cent. f. Bakt., II Abt., 25, 1910, 53.) From Rochelle salts (sodium potassium tartrate). Micrococcus dermatogenes Fuhrmann. (Cent. f. Bakt., II Abt., 17, 1906, 618.) From bottled beer. Micrococcus diffluens Schroeter. (In Cohn, Kryptog. -Flora v. Schlesien, 3, 1, 1886, 144.) From dust, feces, etc. Micrococcus dimorpkus Bucherer. (Planta, Arch. f. wissen. Bot., 1934, 98.) A dimorphic bacterium. He reports it as much like Micrococcus melitensis Bruce and Bacterium fraenkelii Hashi- moto. Micrococcus diphtericus (sic) Cohn. (Micrococcus, Oertel, Deutsch. Arch. f. klin. Med., 8, 1871 ; Cohn, Beitr. z. Biol, d. Pflanzen, i, Heft 2, 1872, 162; Strepto- coccus diphtheriticus Zopf, Die Spalt- pilze, 3 Aufl., 1885, 53.) From throats and nasal passages of diphtheria patients. Micrococcus dissimilis Dyar. (Sec Sattler, Cent. f. Bakt., 5, 1889, 70; Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 353 ; Micro- coccus trachomatis conjunctivae Sattler in Krai, Die gegenwartigen Bestand der Kral'schen Sammlung von Mikroorgan- ismen, 1900, 19.) From trachoma infec- tions. Hucker (loc. cit., 17) considers this a synonym oi Micrococcus caseolyticus Evans. Micrococcus djokjakartensis Zettnow. (Cent. f. Bakt., I Abt., Orig., 75, 1915, 376.) From a sugar factory in Java. Micrococcus doyeni De Toni and Trevi- san. {Micrococcus urinae alhus olearius Doyen, Jour. d. connaiss. medic, No. 14, 1889, 108; De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1076.) From urine. Hucker (loc. cit., 16) states that this species is apparently identical with Micrococcus albus Schroeter. Micrococcus drimophylus Baumgartner. (Baumgartner, Ergebnisse d. ges. Zahn- heilk.. Heft 2, 1910, 729; Abst. in Cent, f. Bakt., I Abt., Ref., 48, 1911, 622.) From the mouth cavitv. Micrococcus eatonii Corbet. (Quart. Jour. Rubber Research Inst. Malaya, 2, 1930, 145.) From the latex of the rubber tree (Hevea hrasiliensis). For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 244. Micrococcus eburneus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 85.) From Camembert cheese. Win- slow and Winslow {loc. cit., 224) state that this species is apparently a synonym of Micrococcus candicans Flixgge. Micrococcus ephestiae Mattes. (Sit- zungsber. d. Gesellsch. z. Beford. d. gesamt. Naturwissensch. zu Marburg, 62, 1927, 406.) From the Mediterranean flour moth {Ephestia kuehniella). Micrococcus epimetheus Corbet {loc. cit., 148). From the latex of the rubber tree {Hevea hrasiliensis). For a de- scription of this species, see Bergey et al.. Manual, 5tli ed., 1939, 256. Micrococcus esterificans Beck. (Arb. kaiserl. Gesundheitsamte, 29, Heft 2, 1905; Abst. in Cent. f. Bakt., II Abt., 19, 1907, 594.) Has a characteristic fruity aroma. From butter. Micrococcus exanthematicus Lewa- scheff. (Deutsch. med. Wochnschr., No. 13 and 34, 1892; Abst. in Cent. f. Bakt., 12, 1892, 635.1 From blood in cases of typhus fever. Motile. Grows anaer- obically. Micrococcus excavatus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 486.) From the stomach contents of a coot {Fulica atra) and a woodpecker (Picus major). Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus exiguus Kern {loc. cit., 470). From the stomach contents of the chaffinch {Fringella coelebs). Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micro- coccus alMis Schroeter; while Hucker (loc. cit., 19) considers it a synonym of Micrococcus Jreudenreichii Guillebeau or of Micrococcus ureae Cohn. Micrococcus expositionis Chester. (No. FAMILY MICROCOCCACEAE 261 34, Conn, Storrs Agr. Exp. Sta. 7th Ann. Rept., 1895, 77; Chester, Man. Determ. Bact., 1901, 92.) From air. Winslow and Winslow {loc. cit. 216) consider this a synonym of Micrococcus flavus Trevi- san; while Hucker (loc. cit., 10) regards it as a synonym of Micrococcus conglom- eratus Migula. Micrococcus expressus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 195.) From a carrot infusion. Produces slime. Hucker {loc. cit., 7) considers this species a synonym of Micrococcus luteus Cohn or of Micrococcus varians Migula. Micrococcus faviformis Migula. (Milchweisser Diplococcus, Bumm, Mi- kroorg. d. gonorrh. Schleimbautkr., II Ausg., 1887, 18; Micrococcus lacteus faviformis Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 182; Neisseria lactea Trevi- san, I generi e le specie delle Batteriacee, Milan, 1889, 32; Migula, Syst. d. Bakt., 2, 1900, 117.) From vaginal and other body secretions. Winslow and Winslow (loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter. Micrococcus feddei Herter. (Micro- coccus XVI, Choukevitch, Ann. Inst. Past., 25, 1911, 354; Botan. Jahresber., 39, II Abt., Heft 4, 1914, 755; Abst. in Cent. f. Bakt., II Abt., 51, 1920, 367.) From the large intestine of a horse. Re- sembles Micrococcus roscidur Matzu- schita. Micrococcus fervitosus Adametz and Wichmann (Bakt. d. Trink- u. Xutz- wasser. Mitt. Oest. Versuchstat. f. Brauerei u. Malzerei, Wien, Heft 1, 1888.) From water. Winslow and Winslow {loc. cit., 205) consider this a synonym of Gaffkya tetragcna Trevisan. Micrococcus fickii Trevisan. {Coccus albus non liquefaciens {Coccus candicans) Fick, Ueber Microorg. in Conjunctival- sack, Wiesbaden, 1887; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33.) From conjunctiva. Micrococcus finlayensis Sternberg. (Rept. on Etiology and Prevention of Yellow Fever, Washington, 1891, 219.) Obtained by Finlay in cultures from the liver and spleen of a yellow-fever cad- aver. Hucker {loc. cit., 11) considers this a synonym of Micrococcus citreus Migula. Micrococcus flaccidifex danai Brown. (Amer. Museum Novit., No. 251, 1927, 5.) Causative agent of wilt disease of monarch butterfly larvae {Danais archip- pus). Considered a sub-species of Gyro- coccus flaccidifex Glaser and Chapman (Science, 36, 1912, 219). Micrococcus flagcllatus Klotz. (Jour. Med. Research, 11 (N.S.6), 1904, 493.) Found in an epizootic among rabbits and white rats. Supposedly flagellated. Micrococcus flavens Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 80.) From Swiss cheese. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus flavescens Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 79.) From Swiss cheese. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavxis Trevisan. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 246. Micrococcus flavidus Henrici. {loc cit., 81; not Micrococcus flavidus Roze, Compt. rend. Acad. Sci., Paris, 122, 1896, 750.) From Swiss and Limburger cheeses. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus flavovirens Migula. {Staphylococcus viridis flavescens Gutt- mann, Arch. f. path. Anat., 107, 1887, 261 ; Staphylococcus viridi-flavescens Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 33; Migula, Syst. d. Bakt., 2, 1900, 124; Micrococcus viridis Chester, Man. Determ. Bact., 1901, 95; Micrococcus viridis -flavescens Winslow and Winslow, Systematic Relationships of the Coccaceae, 1906, 221.) Winslow and Winslow {ibid., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus flavus non liquefaciens 262 MANUAL OF DETERMINATIVE BACTERIOLOGY Amsler. (Amsler, Korrespondenbl. f. Schweizer Aerzte, 1900, No. 9; Abst. in Cent. f. Bakt., I Abt., 29, 1901, 450.) From thermal springs. Micrococcus fluorescens Maggiora. (Giorn. Soc. Ital. d'Igiene, 11, 1889, 352; Abst. in Cent. f. Bakt., 8, 1890, 13.) From the skin of the foot. Micrococcus foetidus Fliigge. (Die Mi- kroorganismen, 2 Aufl., 1886, 172; not Micrococcus foetidus Veillon, Compt. rend. Soc. Biol. Paris, 1893, 867; see Streptococcus foetidus Prevot.) From carious teeth. Micrococcus foetidus Klamann. (All- gem, med. Centralzeitung, 1887, 1344.) Isolated from the posterior nares of man. Winslow and Winslow (loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter. Micrococcus frag His (Dyar) Migula. (Merismopedia fragilis Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 351; Migula, Syst. d. Bakt., 2, 1900, 186.) From dust. Muck- er (loc. cit., 25) states that this species may be identical with M icrococcus roseus Fliigge. Micrococcus franklandioruin Trevisan. (Micrococcus candicans Frankland and Frankland, Phil. Trans. Roy. Soc. Lon- don, 178, B, 1888, 270; not Micrococcus candicans Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 173; Trevisan, I generi o le specie delle Batteriacee, Milan, 1889, 34.) From dust. Micrococcus fulvus Weiss. (Arb. a. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 206; not Micrococcus fulvus Cohn, Beitr. z. biol. d. Pflanz., /, Heft 3, 1875, 181.) From a bean infusion. Micrococcus fuse us Adametz . (Brauner Coccus, Maschek, Jahresb. d. Kommunal- Oberrealsch. zu Leitmeritz, No. 6, 1867, 60; Adametz, Bakt. d. Nutz- u. Trink- wasser, Vienna, 1S88; Micrococcus fuscus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 932.) Hucker (loc. cit., 10) states that this species is prob- ably identical with Micrococcus con- glomeratus Migula. Micrococcus galbanatus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 68.) From water. Winslow and Winslow (loc. cit., 216) consider this a synonym of Micro- coccus flavus Trevisan. Micrococcus gallicidus Burrill. (Amer. Nat., 17, 1883, 320.) From blood of fowls infected with chicken cholera. Micrococcus gelatinogenus Briiutigam. (Pharmaceutische Centralhalle, 32, 1891, 427.) From digitalis infusions. See Micrococcus gummosus Happ. Micrococcus gelatinosus Warrington. (The Lancet, 1, 1888, No. 25; Abst. in Cent. f. Bakt., 4, 1888, 394.) Curdles milk. Micrococcus gelatinosus Issatchenko. (Recherches sur les Microbes de I'Ocean Glacial Arctique, Petrograd, 1914, 232; not Micrococcus gelatinosus Warrington, The Lancet, 1, 1888, No. 25.) From sea water. Micrococcus giganteus lactis Conn. (Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 46.) From milk. Micrococcus gigas Frankland and Frankland. (Philos. Trans. Roy. Soc, London, 178, B, 1888, 268.) From dust. Winslow and Winslow (loc. cit., 216) con- sider this a synonym of Micrococcus flavus Trevisan. Micrococcus gilvus Losski. (Inaug. Diss., Dorpat, 1893, 60.) Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus gingivae Migula. (Mi- crococcus gingivae pyogenes Miller, Die Mikroorganismen d. Mundhohlc, Leipzig, 1889, 216; Migula, Syst. d. Bakt., 2, 1900, 68.) From alveolar pyorrhoea, also from the mouth of a healthy man. Micrococcus gingreardi Renault. (Compt. rend. Acad. Sci., Paris, 120, 1895,217.) Micrococcus glandulosus Weiss. (Arb. bakt. Inst. Karlsruhe, ^, Heft 3, 1902, 201 . ) From an asparagus infusion. Hucker FAMILY MICROCOCCACEAE 263 {loc. cit., 19) regards this speciee as identical with Micrococcus freiidenreichii Guillebeau or with Micrococcus ureae Cohn. Micrococcus globosus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 469.) From the stomach contents of a coot {Fulica atra). Winslow and Wins- lov,' {loc. cit., 224) state that this is appar- ently a synonym of Micrococcus candicans Fliigge. Micrococcus granulatus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 197.) From a malt infusion. Micrococcus granulosus Kern {loc. cit., 483). From the stomach contents of the yellow-hammer {Emheriza citrinella) and the starling {Sturnus vulgaris). Wins- low and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus griseus (Warming) Winter. {Bacterium griseum Warming, Viden- skabelige Meddelelser fra den naturhist. Forening i Kjobenhavn, 1875, 398; Winter, in Rabenhorst, Kryptog. -Flora V. Deutschl., Oesterr. u. d. Schweiz, 2 Aufi., 1, 1884,47.) Micrococcus grossus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 71.) From Camembert cheese. Wins- low and Winslow {loc. cit., 224) state that this is apparently a synonym of Micrococcus candicans Fliigge. Micrococcus gummosus Happ. (Happ, Inaug. Diss., Basel, 1893, 31; not Micro- coccus gummosus Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 189.) From snakeroot and digitalis infusions. Pre- sumably Leuconostoc mesenieroides Van Tieghem. Micrococcus haematodes Zopf. (Mi- crobes de la sueur rouge. Babes, Biol. Centralbl., 2, 1882, No. 8; Zopf, Die Spaltpilze, 3 Aufl., 1885, 60.) The cause of red perspiration. Hucker {loc. cit., 25) states that this may be a synonym of Micrococcus roseus Fliigge. Micrococcus haemorrhagicus (Klein) Migula. {Staphylococcus haemorrhagicus Klein, Cent. f. Bakt., I Abt., 22, 1897, 81; Migula, Syst. d. Bakt., 2, 1900, 88.) Associated with an erythema of the skin resembling anthrax. Winslow and Wins- low {loc. cit., 199) state that this appears to be a synonym of Micrococcus albtis Schroeter. Micrococcus halensis Lehmann and Neumann. {Micrococcus acidi para- lactici liquefaciens halensis Kozai, Ztschr. f. Hyg., 31, 1899, 374; Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 210; Micrococcus acidi paralactici lique- faciens Thiele, Ztschr. f. Hyg., ^6, 1904, 394.) From milk. Hucker {loc. cit., 17) considers this a synonym of Micrococcus caseolyticus Evans. Micrococcus halophilus Bergey et al. (Culture No. 19, Baranik-Pikowsky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al.. Manual, 3rd ed., 1930, 89.) From sea water. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 258. Micrococcus hauseri (Rosenthal) Mi- gula. {Diplococcus hauseri Rosenthal, Inaug. Diss., Berlin, 1893, 26; Migula, loc. cit., 80.) From the oral cavity. Winslow and Winslow {loc. cit., 224) state that this species is apparently identical with Micrococcus candicans Fliigge. Micrococcus helvolus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 77.) From Swiss cheese. Winslow and Winslow {loc. cit., 220) consider this to be identical with Micrococcus luteus Cohn . Micrococcus hibiscus Nakahama. (Jour. Agr. Chem. Soc. Japan, 16, 1940, 345, Eng. Abs., Bull. Agr. Chem. Soc, 16, 1940, 51.) Isolated from retting kenaf {Hibiscus). Micrococcus humidus Migula. (Micro- coccus No. 2, Adametz, Landwirtsch. .lahrb., 18, 1889, 239; Migula, Syst. d. Bakt., 2, 1900, 50.) From Emmenthal cheese. Winslow and Winslow {loc. cit., 224) state that this species is appar- ently identical with Micrococcus candi- cans Fliigge. 264 MANUAL OF DETERMINATIVE BACTERIOLOGY Micrococcus hydrolhermicus Cronquist. (Monatsh. f. prakt. Derm., 36, 1903.) Optimum temperature 41 °C. Micrococcus hymenophagus Renault. (Compt. rend. Acad. Sci., Paris, 120, 1895, 217.) Micrococcus imperatoris Roze. (Compt. rend. Acad. Sci., Paris, 122, 1896, 545.) Isolated from potatoes. Probably from dust. Micrococcus inconspicuus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894,64.) From Swiss cheese. Winslow and Winslow {loc. cit., 224) state that this species is apparently identical with Micrococcus candicans Fliigge. Micrococcus indolicus Christiansen. (Ac. Path. Micr. Scand., 18, 1934, 42; Staphylococcus asacchnrolyticus var. in- dolicus Weinberg, Nativelle and Prevot, Les Microbes Anaerobies, 1937, 1023.) Strict anaerobe. For description see Bergey et al., Manual, 5th ed., 1939, 266. Micrococcus influenzae Migula. (Mi- kroorganismus II, Fischel, Ztschr. f. Heilkunde, 12, 1891; Abst. in Cent. f. Bakt., 9, 1891, 611; Migula, Syst. d. Bakt., 2, 1900, 90.) From the blood of an influenza patient. Winslow and Wins- low {loc. cit., 199) state that this appears to be identical with Micrococcus albus Schroeter. Micrococcus insectoruiu Burrill. (Burrill, Amer. Nat., 17, 1883, 319; Strep- tococcus insectoruni De Toni and Trevi- san, in Saccardo, Sylloge Fungorum, 8, 1889, 1061; Bacillus insectoruiu Smith, 28th Biennial Kept. Kan. State Bd. Agri., 1933, 54.) From the cecal organs of the chinch bug (Blissus leucopterus). Micrococcus intermedius Stark and Scheib. (Jour. Dairy Sci., 19, 1936, 210.) From butter. Micrococcus intertriginis ^eyer. (N.Y. Med. Jour., 70, 1900, 873; Abst. in Cent. f. Bakt., I Abt., 30, 1901, 434.) From a case of erythema intertrigo. Micrococcus iris Henrici. (Arb. bakt. Inst. Karlsruhe, /, Heft 1, 1894, 67.) From Limburger cheese. Winslow and Winslow (loc. cit., 224) state that this species is apparently identical with Micrococcus candicans Fliigge. Micrococcus irregularis Weiss. (Arb. bakt. Inst. Karisruhe, 2, Heft 3, 1902, 184.) From an infusion of beans and carrots. Micrococcus jo7igii Chester. (Staph- ylococcus pyogenes bovis de Jong, Cent, f . Bakt., I Abt., 25, 1899, QA:-, Staphylococcus bovis de Jong, ibid., 67; Chester, Man. Determ. Bact., 1901, 95.) Associated with suppurative processes in cattle. Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus lactericeus Freund. (In- aug. Diss., Erlangen, 1893, 21; Abst. in Cent. f. Bakt., 16, 1894, 640.) From the human mouth. Hucker {loc. cit., 21) regards this as a synonym of Micrococcus candidus Cohn or of Micrococcus epider- midis Hucker. Micrococcus lacteus Henrici. (Arb. bakt. Inst. Karisruhe, 1, Heft 1, 1894, 74.) From cheese. Winslow and Wins- low {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker (loc. cit., 19) considers it a synonym of Micrococcus freudenreichii Guillebeau or of Micro- coccus ureae Cohn. Micrococcus lactis Chester. (No. 44, Conn, Storrs Agr. Exp. Sta., 7th Ann. Rept., 1895, 79; Chester, Man. Determ. Bact., 1901, 90.) From milk. Winslow and Winslow {loc. cit., 224) state that this is apparently a synonym of Micro- coccus candicans Fliigge. Micrococcus lactis II (Hueppe) Scholl. (Quoted from Lohnis, Cent. f. Bakt., II Abt., 18, 1907, 141.) From milk. Micrococcus lactis acidi Krueger. (Cent. f. Bakt., 7, 1890, 494.) From milk. Micrococcus lactis albus Conn, Esten and Stocking. (Storrs Agr. Exp. Sta. 18th Ann. Rept., 1907, 120.) From milk. Hucker {loc. cit., 19) considers this a synonym of Micrococcus freudenreichii FAMILY MICROCOCCACEAE 265 Guillebeau or of Micrococcus ureae Cohn. Micrococcus lactis amari von Freuden- reich. (Bittere Milch Micrococcus, Conn, Cent. f. Bakt., 9, 1891, 653; von Freudenreich, Cent. f. Bakt., II Abt., 13, 1904, 407.) From the udder and bitter cream. Micrococcus lactis aureus Conn, Esten and Stocking (loc. cit., 112). From milk, butter, cheese, stable dust. Hucker {loc. cit., 7 and 12) regards this as a syn- onym of Micrococcus luteus Cohn, of Micrococcus varians Migula or of Micro- coccus aureus Zopf. Micrococcus lactis citreus Conn, Esten and Stocking (loc. cit., 102). From milk. Hucker (loc. cit., 7) considers this species identical with Micrococcus luteus Cohn or with Micrococcus varians Migula. Micrococcus lactis citronus Conn, Esten and Stocking {loc. cit., 117). From slime on Camembert cheese. Hucker {loc. cit., 12) regards this as a synonym of Micrococcus aureus Zopf. Micrococcus lactis flavus Conn, Esten and Stocking {loc. cit., 109). May be identical with Micrococcus aurantiacus Cohn. From milk. Hucker {loc. cit., 7 and 12) states that this may be a syn- onym of Micrococcus luteus Cohn, of Micrococcus varians Migula or of Micro- coccus aureus Zopf. Micrococcus lactis fluorescens Conn, Estenand Stocking {loc. cit., 120). From stable dust. Exhibits a green fluores- cence. Hucker {loc. cit., 18) states that this species is very similar to Micrococcus caseolyticus Evans. Micrococcus lactis gigas Conn, Esten and Stocking {loc. cit., 116). From milk. Hucker {loc. cit., 22) states that this species is probably identical with Micro- coccus candidus Cohn or Micrococcus epidermidis Hucker. Micrococcxis lactis giganteus Conn, Estenand Stocking {loc. cit., 122). From milk. Micrococcus lactis minutissimus Conn, Esten and Stocking {loc. cit., 119). From milk. Hucker (loc. cit., 10) considers this a synonym of Micrococcus conglom- eratus Migula. Micrococcus lactis rosaceus Conn, Esten and Stocking (loc. cit., 109). From milk. Hucker (loc. cit., 26) states that this is probably identical with Micrococcus roseus Fliigge. Micrococcus lactis rugosus Conn, Esten and Stocking (loc. cit., 122). From milk. Micrococcus lactis varians Conn, Esten ;ind Stocking (loc. cit., 121). Com- monly found in milk. May be identical with Micrococcus aureus Zopf. Hucker (loc. cit., 15) states that this may be identical in part with Micrococcus albus Schroeter. Micrococcus lardarius Krassilschtschik. (Memoires Soc. Zool. de France, 9, 1896, 513; Compt. rend. Acad. Sci., Paris, 123, 1896, 428.) From diseased silkworms. Differs from Streptococcus bombycis. Hucker {loc. cit., 22) states that this is probably identical with either Micrococ- cus candidus Cohn or Micrococcus epidermidis Hucker. Micrococcus lembkei Migula. (No. 21, Lembke, Arch. f. Hyg., 29, 1897, 327; Migula, Syst. d. Bakt., 2, 1900, 212.) From feces. Winslow and Winslow (loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn, while Hucker (loc. cit., 11) regards it as probably identi- cal with Micrococcus citreus Schroeter. Micrococcus lentus Migula. (Xo. 22, Lembke, loc. cit., 328; Migula, loc. cit., 209.) From feces. Winslow and Wins- low (loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker (loc. cit., 19) regards it as probably identical with Micrococcus freudenreichii Guillebeau or Micrococcus ureae Cohn. Micrococcus licheniformis Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 482.) From the intestine of the yellow- hammer (Emberiza citrinella) . Winslow and Winslow (loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus lignithum Renault. 266 MANUAL OF DETERMINATIVE BACTERIOLOGY (Compt. rend. Acad. Sci., Paris, 126, 1898, 1828.) A fossil form from lignite. Micrococcus lipolyticus Horowitz- Whissowa. (Ztsehr. f. Untersiuth. d. Lebensmittel, 62, 1931, 602.) A fat splitting micrococcus from salted fish. Micrococcus lipolyticus Stark and Scheib. (Jour. Dairy Sci., 19, 1936, 210; not Micrococcus lipolyticus Horo- witz-Wlassowa, loc. cil.) From butter. Micrococcus liquefaciens Migula. (Mi- crococcus ureae liquefaciens Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 169; Streptococcus nelhebius Trevisan, Igeneri e le specie delle Batteriacee, Milan, 1889, 31; Staphylococcus ureae liquefaciens Lundstrora, Festsclir. d. path. anat. Inst. z. Andenken a.d. 250 jiihrige Beste- hen d. finnland. Univ. z. Helsingfors, 1890; abst. in Cent. f. Bakt., 9, 1891, 672; Migula, Syst. d. Bakt., 2, 1900, 106; Micrococcus aethebius Chester, Man. Determ. Bact., 1901, 77; Urococcus lique- faciens flueggei Mirjuel and Cambier. Traite de Bact., 1902, 627; Micrococcus liquefaciens flueggei Miquel in Lafar, Ilandb. d. tech. :\Iykol., S, 1904-06, 75; Albococcus ureae Kliglcr, Jour. Inf. Dis., 12, 1913, 442; not Micrococcus liquefaciens Bergey et al., Manual, 1st cd., 1923, 67.) From urine. Huckcr (loc. cil., 18 and 19) states that this species may be identi- cal with Micrococcus caseolyticus Evans, Micrococcus freudenreichii Guillebeau oi- Micrococcus ureae Cohn. Micrococcus liquefaciens acidi /and //, Conn. (Storrs Agr. Exp. Sta. 12th Ann. Kept., 1900, 48.) From milk. Hucker (loc. cil., 18) regards this species as very similar to Micrococcus caseolyticus Evans. Micrococcus liquidus Migula. (No. 21, Lembke, Arch. f. Hyg., 26, 1896, 313; Migula, Syst. d. Bakt., 2, 1900, 208.) From feces. Hucker (loc. cit., 15) states that this species appears identical with Micrococcus albus Sehroeter. Micrococcus lobatus Migula. (Siebert, Inaug. Diss., Wiirzburg, 1894, No. 3, 10; Migula, Syst. d. Bakt., 2, 1900, 139.) From the human scalp. Winslow and Winslow (loc. cit., 184) state that this is apparently a synonym of Micrococcus aureus Zopf. Micrococcus loewenbergii Trevisan. (Micrococcus de I'ozene, Lowenberg, Congres des otologistes, 1884 and Union medicale, 1884; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33.) From secretions in ozena. Micrococcus luridus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft, 4, 1897, 480.) From the intestine of the chaffinch (Frin- gilla coelebs). Winslow and Winslow (loc. cit., 220) consider this a synonym of Micrococcus luieus Cohn. Micrococcus luteolus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 82; not .Micrococcus luteolus Irwin and Harrison, Le Lait, 8, 1928, 881.) From cheese. Winslow and Winslow (loc. cit., 216) consider this a synonym of Micro- coccus flavus Trevisan. For a description of Irwin and Harrison's organism, see Bergey et al., Manual, 5th ed., 1939, 249.) Micrococcus luteus var. larvae Haudu- ro3" et a\ . (M icrococcus luteus -liquefaciens var. larvae Toumanoff, Bull. Soc. Centr. de Med. Veter., 80, 1927, 367; Hauduroy et al.. Diet. d. Bact. Path., 1937, 277.) From foulbrood of bees. Pathogenic. Micrococcus lutosus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 489.) From the stomach contents of the chaf- finch (Fringilla coelebs). Winslow and Winslow (loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus lysodeikticus Fleming. (Proc. Roy. Soc. London, Ser. B, 93, 1922, 306.) Non-pathogenic. Micrococcus lyssae (Rivolta) Trevisan. (Cocco-baclerium lyssae Rivolta, 1886; Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 33.) Spore-bearer. Micrococcus madidus Migula. (No. 19, Lembke, Arch, f . Hyg., 26, 1896, 311 ; Migula, Syst. d. Bakt., 2, 1900, 207.) From feces. Winslow and Winslow (loc. cit., 184) state that this is apparently a synonym of M icrococcus aureus Aligula ; FAMILY MICROCOCCACEAE 26'; while Hucker {loc. cit., 15) regards it as identical with Micrococcus aJbris Schroetcr. Micrococcus inagnus (Miller) Trevisan. {Jodococcus magnus Miller, Deutsche mod. Wochnschr., Xo. 30, 1888 ; Trevisan, I generi c le specie dellc Batteriacee, Milan, 1889, 33; not Micrococcus magnus Chester, Man. Determ. Bact., 1901, 85 ( Diplococcus magnus Rosenthal, Cent. f. Bakt., 25, 1899, 1); not Micrococcus magnus Stark and Scheib, .lour. Dairy Sci., 19, 1936, 210.) Micrococcus major Dc Toni and Trevi- san. (Micrococcus urinae major Doyen, Jour. d. connaiss. medic, Xo. 14, 1889, 108; De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1076.) From urine. Micrococcus major Eckstein. (Ztschr. f. Forst- u. Jagdwesen. 26. 1894, 18; not Micrococcus major DeToni and Tre- visan, in Saccardo, Sylloge Fungorum, 8, 1889, 1076.) Isolated from the larvae of the nun moth (Lymantria monacha) and Hypono77ieuta sp. Micrococcus manfredii Trevisan. (Mi- crococcus der progressiven Lymphome im Tierorper, Manfredi, Fortschr. d. Med., 1886, 713; Trevisan, I generi e Ic specie delle Batteriacee, Milan, 1889, 33; Streptococcus manfredii DeToni and Trevisan, in Saccardo, Sylloge Fungo- rum, 8, 1889, 1056; Micrococcus canus Migula, Syst. d. Bakt., 2, 1900, 63.) From sputum. Winslow and Winslow {loc. cit., 206) regard this as a synonym of Micrococcus candidus Colin or of Gaffkya tetragena Trevisan. Micrococcus malolacticus Scifert. (Ztschr. f . d. landwirtsch. Versuchswesen in Oesterreich, 1903, 567; Abst. in Cent, f. Bakt., II Abt., 10, 1903, 664.) From wine. Hucker (loc. cit., 7) considers this a synonym of Micrococcus luteus Cohn or of Micrococcus varians ISIigula. Micrococcus mammitis Hutchens. (Hutchens in Besson, Pract. Bact. Microbiol, and Serum Therapy, Trans, of 5th ed., 1913, 615; not Streptococcus mammitis hovis Hutchens, ibid., 613.) From gangrenous mammitis of milking ewes. This is le microcoque de I'ar- aignee or de la mammite gangreneuse, Xocard, Ann. Inst. Past., 1, 1887, 417. Micrococcus marginaius Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 11 .) From meat extract. Micrococcus marinus Issatchenko. (Recherches sur les Microbes de I'Ocean Glacial Arcticjue, Pctrograd, 1914, 147.) From sea water. Micrococcus mastitidis Holland. (Jour. Bact.. 5, 1920, 224.) Nomen nudum. Micrococcus mastohius Trevisan. (Microcoque de la mammite gangreneuse des brebis laitieres, Xocard, Ann. Inst. Past., 1, 1887, 417; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33; Micrococcus gangr. avium, quoted from Freire, Rio de Janeiro, 1898; Abst. in Cent. f. Bakt., I Abt., 26, 1899, 841; Micrococcus ovis Migula-, Syst. d. Bakt., 2, 1900, 90; Micrococcus mastilis Chester, Man. Determ. Bact., 1901, 76; Micrococ- cus mastidis gangraenosae ovis Pfeiler, Ztschr. f. Infektionskr., parasit. Krankh. u. Hyg. d. Haustiere, 4, 1908, 132.) Said to be the cause of gangrenous mastitis in sheep. This species appears to be Micrococcus pyogenes Migula. Micrococcus melanocyclus Merker. (Cent. f. Bakt., II Abt., 41, 1911, 589.) See Actinomyces melanocyclus Krainsky. Micrococcus melanoglossophorus Spe- gazzini. (Fung. Arg. Pug., 4, 18--, 316.) From the epithelium of the tongue. Micrococcus meldensis Roger. (Ber. An. de Soc. d'Agric. de Meau.x, 1898.) Micrococcus mclleus grandinis Harri- son. (Bot. Gazette, 26, 1898, 211.) Micrococcus memelensis Leichmann. (Coccus, Leichmann, Cent. f. Bakt., II Abt., 2, 1896, 780; Leichmann, in Koch, Jahresber., 12, 1901, 254; Micro- coccus acidi laevolactici Weigmann, in Lafar, Handb. d. techn. Mykol., 2, 1905, 62.) From milk. Micrococcus minimus ^yeiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 268 MANUAL OF DETERMINATIVE BACTERIOLOGY 188; not Micrococcus minimus Bergey et al., Manual, 1st ed., 1923, 69.) From a bean infusion. Hucker (loc. cit., 7) considers this a synonym of either Micro- coccus luteus Cohn or Micrococcus varians Migula. Micrococcus minutissimus Issatclienkij. (Recherches sur les Microbes de I'Ocean Glacial Artique, Petrograd, 1914, 146.) From sea water. Micrococcus mirificus (Rabenhorst) Trevisan. {Palmella miriUca Raben- horst, Hedwigia, 1867, 115, and Flor. Europ. Algar., 3, 1856, 35; Trevisan, Rendic. R. 1st. Lombardo, 12, 1879.) Micrococcus mollis (Dyar) Migula. (Merismopedia mollis Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 352; Migula, Syst. d. Bakt., 2, 1900, 161; Aurococcus mollis Winslow and Rogers, Science, 21, 1905, 669; Staphylococcus mollis Holland, Jour. Bact., 5, 1920, 225.) From air. A cause of boils in the tropics, according to Castel- lani and Chalmers (Man. Trop. Med., 3rd ed., 1919, 931). Hucker {loc. cit., 12) states that this species is apparently identical with Micrococcus aureus Zopf. Micrococcus (Diplococcus) morrhuae Klebahn. (Mitteil. Inst. Allgm. Botan. Hamburg, 4, 1919, 11-69; Abst. in Cent. f. Bakt., II Abt., S£, 1921, 123.) Halo- philic. Associated with spoilage of salted fish. Micrococcus mucilagineus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 191.) From bean infusions. Hucker (loc. cit., 11) states that this is probably a synonym of Micrococcus citreus Migula. Micrococcus mucilaginosus Migula. (Micrococcus der schleimigen Milch, Ratz, Arch. f. Tierheilkunde, 12, Heft 1 and 2, 1890; Migula, Syst. d. Bakt., 2, 1900, 119; not M icrococcus mucilaginosus Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 205.) From slimy milk. Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker {loc. cit., 18) considers it a syn- onym in part of Micrococcus caseolyticus Evans. Micrococcus mucofaciens Thoni and Thaysen. (Cent. f. Bakt., II Abt., 36, 1913, 359; not Micrococcus mucofaciens Pribram, Klassifikation der Schizomy- ceten, 1933, 42.) From milk. Hucker {loc. cit., 9) considers this a synonym of Micrococcus flavus Trevisan. For a de- scription of this species, see Bergey et al.. Manual, 5th ed., 1939, 245. Micrococcus myceticus Castellani. (Proc. Soc. Exp. Biol, and Med., 25, 1928, 535-536.) From gummy lesions. Micrococcus mycodermatus Holland. (Jour. Bact., 5, 1920, 224.) Micrococcus nacreaceus Migula. (Perl- rauttergliinzender Diplococcus, Tataroff, Inaug. Diss., Dorpat, 1891, 70; Migula, Syst. d. Bakt., 2, 1900, 62.) Winslow and Winslow {loc. cit., 224) state that this is apparently a synonym of Micro- coccus candicans Fliigge. Micrococcus neoformans Doyen. (Doy- en, Le Micrococcus neoformans et les n^oplasmes, Paris, 1903.) From can- cerous tissue. Shown by Andrewes and Gordon (35th Ann. Rept. Local Govt. Board, London, 1905-06, 553) to be iden- tical with Micrococcus epidermidis albus Welch. Micrococcus neurotomae Paillot. (Compt. rend. Acad. Sci., Paris, 178, 1924, 246.) Gram-negative. From the larvae of Neurotoma nemoralis. Micrococcus neuvillei Trevisan. (Mi- crococcus G, Malapert-Neuville, 1887; Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889,34.) From mineral water. Micrococcus nigrescens Castellani. (Brit. Jour, of Dermatology, 23, 1911, 341 ; Nigrococcus nigrescens Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 2103.) Produces a black pigment. Found in the black variety of trichomy- cosis axillaris, a tropical disease. Micrococcus nigrofaciens Northrup. (Mich. Agr. Exp. Sta. Tech. Bull. No. 18, 1914, 12; also in Cent. f. Bakt., II Abt., FAMILY MICROCpCCACEAE 269 41, 1914, 326.) From diseased larvae of the June beetle {Lachnosterna sp.) and other insects. ^ Micrococcus nitidus Kern. (Arb. bakt . Inst. Karlsruhe, 1, Heft 4, 1897, 476.) From the stomach and intestine of birds. Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker {loc. cit., 19) regards it as a syn- onym of Micrococcus freudenreichii Guil- lebeau or of Micrococcus ureae Cohn. Micrococcus nitrificans Bergey et al. (Micrococcus 6, Rubentschick, Cent. f. Bakt., II Abt., 72, 1927, 125; Bergey et al.. Manual, 3rd ed., 1930, 88; not Micro- coccus nitrificans van Tieghem, Traite de Botanique, Paris, 1883.) From sew- age filter beds. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 257. Micrococcus nivalis Chester. (No. 47, Conn, Storrs Agr. Exp. Sta. 7th Ann. Rept., 1895, 80; Chester, Man. Determ. Bact., 1901, 90.) From dust. Winslow and Winslow (loc. cit., 224) state that this is apparently a sj^nonym of Micro- coccus candicans Fltigge. Micrococcus niveus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 66.) From Swiss cheese. Winslow and Winslow {loc. cit., 224) state that this is apparently a synonym of Micrococcus candicans Fliigge. Micrococcus nonfermentans Steinhaus. (Jour. Bact., 42, 1941, 779.) From the alimentary tract of the lyreman cicada {Tibicen linnei) and of an unidentified damsel fly {Coenagrionidae) . Micrococcus nubilis Migula. (Coccus B, Foutin, Bakt. Untersuch. von Hagel, Wratsch, No. 49 and 50, 1889; see Cent, f. Bakt., 7, 1890, 373; Migula, Syst. d. Bakt., 2, 1900, 60 ; M icrococcus beta Ches- ter, Man. Determ. Bact., 1901, 87.) Iso- lated from hail. Winslow and Winslow {loc. cit., 205) consider this to be a syn- onym of Micrococcus candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcus nuclei Roze. (Compt. rend. Acad. Sci., Paris, 122, 1896, 544.) From potatoes. Micrococcus obscoenus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 473.) From the stomach contents of a crow {Corvus corone). Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker (loc. cit., 19) considers it a synonym of Micrococcus freudenreichii Guillebeau or of Micro- coccus ureae Cohn. Micrococcus ochraceus Rosenthal. (In- aug. Diss., Erlangen, 1893, 22; Abst. in Cent. f. Bakt., 16, 1894, 1024; not Micro- coccus ochraceus Hansgirg, Oestr. Bot. Ztschr., 1885, No. 4.) From the oral cavity. Winslow and Winslow {loc. cit., 220) consider this a synonym of Micro- coccus luteus Migula. For a description of this species see Bergey et al., Manual, 5th ed., 1939, 242. Micrococcus ochroleiicus Prove. (Prove, Beitr. z. Biol. d. Pflanz., 4, Heft 3, 1887, 409; Streptococcus ochroleucus Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 31; Planococcus ochroleucus Migula, Syst. d. Bakt., 2, 1900, 272.) Fromi urine. Motile. Micrococcus odoratus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 73.) From cheese. Winslow and Wins- low {loc. cit., 224) state that this is appar- ently a synonym of Micrococcus candicans Flugge. Micrococcus odorus Henrici {loc. cit., 72). From cheese. Winslow and Wins- low {loc. cit., 224) state that this species is apparently a synonym of Micrococcus candicans FHigge. Micrococcus olearius DeToni and Trevisan. {Micrococcus urinae flavus oZear jMsDoyen, Jour. d. connaiss. Medic, No. 14, 1889, 108; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1077.) From urine. Hucker {loc. cit., 12) considers this species a synonym of Micrococcus aureus Zopf. Micrococcus olens Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 87.) 270 MANUAL OF DETERMINATIVE BACTERIOLOGY From Swiss cheese. Winslow and Wins- low {loc. cit., 216) consider this a syn- onym of Micrococcus flavus Trevisan. Micrococcus opalescens DeToni and Trevisan. (Micrococcus albus II, Mag- giora, Giorn. Soc. Ital. d'Igiene, 11, 1889, 351; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1078.) Micrococcus orbicularis Chester. {Mi- crococcus orbicularis flavus Ravenel, Mem. Nat. Acad. Sci.,8, 1896, 8; Chester, Man. Determ. Bact., 1901, 101.) From soil. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micro- coccus flavus Trevisan; while Hucker {loc. cit., 10) regards it as a synonym of Micrococcus conglomeratus Migula. Micrococcus orbiculatus Wright. (Mem. Nat. Acad. Sci., 7, 1895, 432.) From Schuylkill River water. Winslow and Winslow {loc. cit., 220) consider this a synonym of Micrococcus luteus Cohn. Micrococcus ovalis Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 500; not Micrococcus ovalis Escherich, Die Darmbakterien des Sauglings, Stuttgart, 1886, 90.) From the stomach contents of the rock dove {Columba livia). Hucker (loc. cit., 9) regards this as a synonym of Micrococcus flavtis Trevisan. Micrococcus fallens Henrici. (Arb. bakt. Inst. Karlsruhe, /, Heft 1, 1894, 61.) From cheese. Winslow and Wins- low {loc. cit., 205) consider this a syn- onym of Micrococcus candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcus -pallidus Henrici {loc. cit., 62). From cheese. Hucker {loc. cit., 7) regards this species as identical with either Micrococcus luteus Cohn or Micrococcus varians Migula. Micrococcus pannosus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 466.) From the stomach contents of the rock dove {Columba livia) and the in- testine of another dove {Columba oenas). Winslow and Winslow {loc. cit., 224) state that this is apparently a synonym of Micrococcus candicans Fliigge. Micrococcus paraffinae Sohngen. (Cent. f. Bakt., II Abt., S7, 1913, 595.) From garden earth. Micrococcus parotitidis Korentschew- sky. (Cent. f. Bakt., I Abt., Orig., U, 1907, 402.) Isolated from cases of paro- titis epidemica. Micrococcus parvus (Miller) Trevisan. {Jodococcus parvus Miller, Deutsche med. Wchnschr., No. 30, 1888; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33.) Micrococcus parvus Migula. (No. 14, Lembke, Arch. f. Hyg., 26, 1896, 309; Migula, Syst. d. Bakt., 2, 1900, 200.) From feces- Winslow and Winslow {loc. cit., 224) state that this species is appar- ently a synonym of Micrococcus candi- cans Fliigge. Micrococcus pasteuri Trevisan. (Mi- crobe pyogene de I'eau de Seine, Pasteur, 1877; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 34; not Micro- coccus pasteuri Sternberg, Trans. Pathol. Soc. of Philadelphia, 12, 1885, 162.) From water. Micrococcus pellucidus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 468; not Micrococcus pellucidus Roze, Compt. rend. Acad. Sci., Paris, 122, 1896, 1012.) From the intestine of a spar- row {Passer montanus) . Hucker {loc. cit., 23) regards this as a synonym of Micro- coccus candidus Cohn or of Micrococcus epidermidis Hucker. Micrococcus pemphigiMlgula.. (Diplo- coccus des Pemphigus acutus, Demme, Verhandl. d. Kongr. f. innere Med., Wiesbaden, 1886, 336; Diplococcus pem- phigi acuti Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 173; Migula, Syst. d. Bakt., 2, 1900, V and 191; Micro- coccus demmei Chester, Man. Determ. Bact., 1901, 74.) Isolated from bullae in a case of pemphigus acutus. Micrococcus pemphigicontagiosi Castel- lani and Chalmers. {Micrococcus pem- phigi contagiosa Clegg and Wherry, Jour. Inf. Dis., 3, 1906, 171; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 931 .) From bullae in a case of pem- FAMILY MICROCOCCACEAE 271 phigus contagiosa. This may be a synonym of Micrococcus pemphigi neona- torum, see below. Micrococcus pemphigineonatorum Cas- tellani and Chalmers. {Micrococcus pemphigi neonatorum Almquist, Ztsclir. f. Hyg., 10, 1891, 253; Staphylococcus pemphigi neonatorum Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 173; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 931.) Found in bullae in a case of pemphigus neonatorum. This may be Micrococcus mollis, accord- ing to Castellani and Chalmers {loc. cit.). Falls (Jour. Inf. Dis., 20, 1917, 97) identi- fies this and the previous organism as Micrococcus pyogenes var. aureus Zopf. Micrococcus percitreus Bergey et al. (Manual, 1st ed., 1923, 63.) From air and water. Hucker {loc. cit., 10) con- siders this a synonym of Micrococcus conglomeratus Migula. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 248. Micrococcus perUavus Bergey et al. (Manual, 1st ed., 1923, 62.) From air and water. Hucker {loc. cit., 12) re- gards this as a synonym of Micrococcus aureus Zopf. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 247. Micrococcus persicus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 499.) From the intestine of a dove {Co- lumba oenas). Hucker {loc. cit., 25) states that this may be identical with Micrococcus roseus Fliigge. Micrococcus petechialis Trevisan. (Mi- crococco del dermotifo, Bareggi, 1886; Trevisan, I generi e le specie delle Bat- teriacee, Milan, 1889, 33.) Micrococcus petilus Trevisan. (Mi- crococcus der Pyaemie bei Kaninchen, Koch, tjber d. Aetiolog. d. Wundinfec- tionskr., Leipzig, 1878 ; Micrococcus pyae- miae cuniculorum Schroeter, in Cohn, Kryptogam. Flora v. Schlesien, 3, 1, 1886, 148; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, ZZ; Micro- coccus cuniculorum Migula, Syst. d. Bakt., 2, 1900, 192.) From rabbits. Micrococcus peiroZez Renault. (Compt. rend. Acad. Sci., Paris, m, 1897, 1315.) A fossil form from oil bearing rocks. Micrococcus pieridis Burrill. (Quoted from Chittenden, U. S. Dept. Agr., Farmers' Bull. No. 1461, 1926, 6.) From larvae of the cabbage butterfly {Pieris rapae). Micrococcus pikowskyi Bergey et al. (Culture No. 22, Baranik-Pikowsky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al.. Manual, 3rd ed., 1930, 78.) From sea water. For a description of this species, see Bergey et al., Manual, 5th ed., 1939,242. Micrococcus piliformis Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 194.) From a bean infusion. Hucker {loc. cit., 7) considers this a synonym of Micrococcus luteus Cohn or of Micrococ- cus varians Migula. Micrococcus piltonensis Gray and Thornton. (Cent. f. Bakt., II Abt., 73, 1928, 81.) From manure and soil. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 259. Micrococcus pituitoparus (Hohl) Bu- chanan and Hammer. {Karphococcus {Carphococcus) pituitoparus Hohl, Jahrb. d. Schweiz, 22, 1906, 439; Diplococcus viscosus Sato, Cent. f. Bakt., II Abt., 19, 1907, 27; Buchanan and Hammer, Iowa Agr. Exp. Sta. Res. Bull. 22, 1915, 285.) From slimy milk and from straw. Hucker {loc. cit., 23) states that this species is probably identical with Micro- coccus candidus Cohn or with Micrococcus epidermidis Hucker. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 243. Micrococcus plumosus Eisenberg. (Brautigam, Inaug. Diss., Leipzig, 1886, 18; Federiger Micrococcus, Adametz, Mitteil. d. Oesterr. Versuchssta. f. Brau- erei u. Malzerei, Wien, Heft 1, 1888; Eisenberg, Bakt. Diag., 3 Aufl., 1891, 56.) From feces of cattle and from water. Winslow and Winslow {loc. cit., 272 MANUAL OF DETERMINATIVE BACTERIOLOGY 220) consider this a synonym of Micru- coccus luteus Cohn; while Hucker {loc. cit., 22 and 23) regards it as probably identical with Micrococcus candidus Cohn or Micrococcus epidermidis Hucker. Micrococcus polypus Migula. (Syst. d. Bakt., 2, 1900, 79.) From air. Hucker {loc. cit., 23) states that this species is probably identical with Micro- coccus candidus Cohn or Micrococcus epidermidis Hucker. Micrococcus popxdi Delacroix. (Bui. Mens. Off. Renseig. Agr., Paris, 5, 1906, 1349 and Ann. Inst. Nat. Agron., 2 Ser., S, 1906, 353.) Parasitic on poplar trees {Popuhis spp.). Micrococcus porcellorum Trevisan. (Micrococcus bei Hepatitis enzootica porcellorum, Nonewitsch, Cent. f. Bakt., 3, 1888, 233 ; Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 33.) From an infected liver. Micrococcus progrediens Schroeter. (Micrococcus der progressiven Abscess- bildung bei Kaninchen, Koch, Uber d. Aetiolog. d. Wundinfectionskrankheiten, Leipzig, 1878; Schroeter, in Cohn, Kryp- togam. -Flora v. Schlesien, 3, 1, 1886, 148; Micrococcus haernatosaprus Trevi- san, I generi e le specie delle Batteriacee, Milan, 1889, 33.) From the blood of diseased rabbits. Micrococcus psalteri Buemann. (Cent. f. Bakt., I Abt., Orig., 71, 1913, 308.) From the third stomach of cattle. Micrococcus pseudocyaneus Schroeter. (Kryptogam.- Flora v. Schlesien, 3, 1, 1886, 145.) A synonym of Micrococcus cyaneus Cohn according to Migula, Syst. d. Bakt.,^, 1900, 188. Micrococcus pseudoinfluenzae Migula. (Microorganismus I, Fischel, Ztschr. f. Heilkunde, 12, 1891; See Cent. f. Bakt., 9, 1891, 611; Migula, Syst. d. Bakt., 2, 1900, 86.) From the blood of an influ- enza patient. Hucker (loc. cil., 23) considers this a synonym of Micrococcus candidus Cohn or of Micrococcus epi- dermidis Hucker. Micrococcus pulcher Glage. (Ztschr. f. Fleisch- u. Milchhyg., 10, 1900, 146; not Micrococcus pulcher Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 182.) From coating on surface of wurst and similar meat products. Micrococcus pultiformis Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 474.) From stomach contents of the yellow-hammer (Emberiza citrinella) and starling {Sturnus vulgaris) and from the intestine of the woodpecker {Picus ma- jor). Winslow and Winslow {loc. cit., 199) state that this appears to be a syn- onym of Micrococcus alb us Schroeter; while Hucker {loc. cit., 19) regards it as probably identical with Micrococcus freudenreichii Guillebeau or with Micro- coccus ureae Cohn. Micrococcus punctatus Migula. (No. 18, Lembke, Arch. f. Hyg., 29, 1897, 325; Migula, Syst. d. Bakt., 2, 1900, 213.) From feces. Winslow and Winslow {loc. cit., 199) state that this species appears to be a synonym of Micrococcus albus Schroeter. Micrococcus purpurifaciens Lehmann and Neumann. (Micrococcus, Dudt- schenko. Cent, f . Bakt., II Abt., 4^, 1915, 529; Lehmann and Neumann, Bakt. Diag., 6 Aufl., 2, 1920, 755.) From ice. Produces a purple pigment in alkaline gelatin media. Micrococcus p^istulatus Henneberg. (Cent. f. Bakt., II Abt., 65, 1922, 251.) From the human intestine. Micrococcus putridus Tilanus. (Munch. med.Wchnschr., 34, 1887, 310.) From gelatin, agar, etc., containing iodi- form. Micrococcus pygmaexis Henneberg {loc. cit., 252). From the human intestine. Micrococcus pyocyaneus Francisco. (Revista Valenciana de Ciencias Medicas, 1914, 2; Abst. in Cent. f. Bakt., I Abt., Ref., 63, 1915, 44; not Micrococcus pyo- cyaneus Gessard, Thesis, Paris, 1882.) From an acne pustule. Micrococcus pyosepticus (Hericourt and Richet) Solowjew. {Staphylococcus pyo- septicus Hericourt and Richet, Compt. FAMILY MICROCOCCACEAE 273 rend. Acad. Sci., Paris, 107, 1888, 691; Solowjew, Abst. in Cent. f. Bakt., I Abt., 18, 1895, 60.) From an abscess in a dog and from dust. Regarded as identi- cal with Micrococcus albus Schroeter. Micrococcus quadrigeminus Klebs. {Staphylococcus quadrigeminus Vanselow and Czaplewski, Cent. f. Bakt., I Abt., 25, 1899, 143 ; see Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 174.) Closely related to Micrococcus albus Schroeter. Micrococcus quaternus Migula. (Sie- bert, Inaug. Diss., Wiirzburg, No. I, 1894, 7; Migula, Syst. d. Bakt., 2, 1900, 92.) Winslow and Winslow {loc. cit., 199) state that this appears to be a syn- onym of Micrococcus albus Schroeter. Micrococcus radiatus Fliigge. (Die Mikroorganismen, 2 Aufl., 1886, 176; Streptococcus radiatus Crookshank, Man. of Bact., 3rd ed., 1890, 256; not Micro- coccus radiatus Kern, see below.) From dust and water. Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker {loc. cit., 18) considers it a synonym of Micrococcus caseolyticus Evans. Micrococcus radiatus Kern. (Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 471 ; Micrococcus radiosus Migula, Syst. d. Bakt., 2, 1900, 114.) From the stomach contents of the starling {Sturnus vulgaris). Winslow and Winslow (loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter. Micrococcus reessii Rosenthal. (Inaug. Diss., Berlin, 1893, 19; Abst. in Cent. f. Bakt., 16, 1894, 1024.) From the oral cavity. Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter. Micrococcus regular is Weiss. (Arb. bakt. Inst. Karlsruhe,^, Heft 3, 1902, 183.) From bean infusions. Hucker {loc. cit., 7) considers this a synonym of Micrococ- cus luteus Cohn or Micrococcus varians Migula. Micrococcus resinaceus Kern. (Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1897, 487.) From the stomach contents of the starling {Sturnus vulgaris) and from the intestine of a sparrow {Passer montanus). Winslow and Winslow {loc. cit., 220) re- gard this as a synonym of Micrococcus luteus Cohn. Micrococcus rhenanus Migula. (Neuer Mikrococcus aus Rheinwasser, Burri Arch. f. Hyg., 19, 1893, 34; Migula, Syst d. Bakt., 2, 1900, 109; Micrococcus rhen Chester, ]\Ian. Determ. Bact., 1901, 82 Albococcus rhejianus Winslow and Rogers, Jour. Inf. Dis., 3, 1906, 544.) From Rhine River water. Winslow and Wins- low {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter; while Hucker {loc. cit., 18) considers it a synonym of Micrococcus caseolyticus Evans. Micrococcus ridleyi Corbet. (Quart. Jour. Rubber Research Inst., Malaya, 2, 1930, 146.) From the latex of the rubber tree {Hevea brasiliensis) . For a de- scription of this species, see Bergey et al., Manual, 5th ed., 1939, 244. Micrococcus rosaceus Frankland and Frankland. (Trans. Roy. Society, London, 178, B. 188, 269; Rhodococcus rosaceus Holland, Jour. Bact., 5, 1920, 225.) From air. Hucker {loc. cit., 25) states that this species may be identical with Micrococcus roseus Fliigge. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 252. Micrococcus rosaceus lactis Conn. (Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 34; Micrococcus lactis rosaceus Conn, Esten and Stocking, Storrs Agr. Exp. Sta. Rept. for 1906, 108.) From milk. Micrococcus roscidus Migula. (Micro- coccus No. I, Adametz, Landwirtsch. .lahrb., 18, 1889, 238; Migula, Syst. d. Bakt., 2, 1900, 68.) From Emmenthal cheese. Winslow and Winslow {loc. cit., 224) state that this is apparently a synonymofMi'croccccwscandiccns Fliigge. Micrococcus roseo-persicinus Migula. 274 MANUAL OF DETERMINATIVE BACTERIOLOGY (Rote Kokken von Van Ermengem, Schneider, Arb. bakt. Inst. Karlsruhe, 1, Heft 2, 1894, 216; Migula, Syst. d. Bakt., 2, 1900, 184.) Micrococcus rosettaceus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, I Reihe, 1890, 72.) From water. Winslow and Winslow {loc. cit., 224) state that this is apparently a synonym of Micrococcus candicans Flugge. Micrococcus 7-oseus Maggiora. (Giorn. Soc. Ital. d'Igiene, 11, 1889, 356; not Micrococcus roseus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 183; not Mi- crococcus roseus Gruber, Cent. f. Bakt., II Abt., 22, 1909, 408.) Micrococcus rubellus Migula. (Syst. d. Bakt., 2, 1900, 169.) Source not given. Hucker {loc. cit., 27) regards this as identical with Micrococcus cinnabareus Flugge. Micrococcus rubescens Migula. (Xo. 20, Lembke, Arch. f. Hyg., 26, 1896, 312; Migula, Syst. d. Bakt., 2, 1900, 208; not Micrococcus rubescens Chester, see Micrococcus subroseus below.) Frorh feces. Hucker {loc. cit., 27) regards this species as identical with Micrococcus cinnabareus Flugge. Micrococcus rubidus lactis Conn. (Conn, Storrs Agr. Exp. Sta. 12th Ann. Rept., 1900, 34; Micrococcus lactis rubi- dus Conn, Esten and Stocking, Storrs Agr. Exp. Sta. 18th Ann. Rept., 1907, 117.) From milk. Resembles Micro- coccus cinnabareus Fliigge. Hucker {loc. cit., 25) thinks this species may be iden- tical with Micrococcus roseus Fliigge. Micrococcus rubigerwsus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 492.) From the stomach contents of a dove {Columba oenas). Hucker {loc. cit., 25) states that this species may be identical with Micrococcus roseus Fliigge. Micrococcus rubiginosus Passer, and Beltr. (Fung. Sicil., 18—, no. 35; quoted from DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1082.) Micrococcus rugatus Migula. {Micro- coccus endocarditidis rugatus Weichsel- baum, Beitr. z. path. Anat. u. z. allgm. Pathol., 4, 1889, 164; Migula, Syst. d. Bakt., 2, 1900, 190; Micrococcus endo- carditis Chester, Man. Determ. Bact., 1901, 74.) From ulcerative endocarditis. Winslow and Winslow {loc. cit., 205) con- sider this a synonym of Micrococcus can- didus Cohn or of Gaffkya tetragena Tre- visan; while Hucker {loc. cit., 15) regards it as a synonym of Micrococcus albus Schroeter. Micrococcus rugosus Chester. (No. 2, Conn, Storrs Agr. Exp. Sta. 6th Ann. Rept., 1894, 50; Chester, Man'. Determ. Bact., 1901, 101.) From milk and ripened cream. Winslow and Winslow {loc. cit., 216) consider this a synonym of Micrococcus flavus Trevisan. Micrococcus ruminantium Henneberg. (Cent. f. Bakt., II Abt., 55, 1922, 252.) From the human intestine. Micrococcus rushrnorei Brown. (Amer. Museum Novit., No. 251, 1927, 4.) Iso- lated from a fly {Lucilia sericata) which was infected with Bacillus lutzae. Micrococcus saccatus Migula. {Micro- coccus albus liquefaciens von Besser, Beitr. z. i)ath. Anat., 6, 1889, 46; Micro- coccus liquefaciens albus, see Cent. f. Bakt., 7, 1890, 152; Migula, Syst. d. Bakt., 2, 1900, 117; Micrococcus lique- faciens Chester, Man. Determ. Bact., 1901, 78; not Micrococcus liquefaciens Holland, Jour. Bact., 5, 1920, 224; Mi- crococcus alvi Chester, loc. cit., 81.)From the nasal mucous membrane. Winslow and Winslow {loc. cit., 199) state that this is apparently a synonym of Micro- coccus albus Schroeter; while Hucker {loc. cit., 19) regards it as probably identi- cal with Micrococcus freudenreichii Guille- beau or with Micrococcus ureae Cohn. For a description of this species, see Bergey et al.. Manual, 5th ed., 1939, 254. Micrococcus salivalis septicus, quoted from Wigura, see Cent. f. Bakt., I Abt., 17, 1895, 899. From the human skin. Micrococcus sarcinoides Migula. (Syst. d. Bakt., 2, 1900, 168.) Hucker FAMILY MICKOCOCCACEAE 97; (loc. cit., 27) considers this identical witli Micrococcus cinnabareus Fliigge. Micrococcus scariosus Migula. (Sie- bert, Inaug. Diss., Wiirzburg, No. II, 1894, 9; Migula, Syst. d. Bakt., 2, 1900, 91.) From a hairbrush. Winslow and Winslow {loc. cit., 199) state that this appears to be a synonym of Micrococcus albus Schroeter. Micrococcus scarlatiriosus Trevisan. (Trevisan, Batteri itaiiani, 1879, 19; Streptococcus rubiginosus Edington, Brit. Med. Jour., 1, 1887, 1265; Perroncitoa scarlatinosa Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 29.) From a scarlet fever patient. Micrococcus scarlatinus ^ligula. (Syst. d. Bakt., 2, 1900, 173.) From feces. Micrococcus selenicus Brenner. (Jahrb. f. wissensch. Botan., 57, 1916, 85; Abst. in Cent. f. Bakt., II Abt., 48, 1918, 431.) From mud. Micrococcus sensibilis Zettnow. (Cent. f. Bakt., I Abt., Orig., 77, 1915, 216.) From dust. Hucker (?oc. cz'i., 19) considers this a synonj'm of Micrococcus freudenreichii Guillebeau or of Micrococ- cus ureae Cohn. For a description of this species, see Bergey et al., Manual, 5th ed., 1939, 248. Micrococcus septicus (Klebs) Cohn. {Microsporon septicum Klebs, Die Ur- sachen der infectiosen Wundkrankheiten, 1871; and Zur path. Anat. d. Schuss- wunden, 1872; Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 2, 1872, 164.) From pus. Micrococcus serophilus Costa. (Compt. rend. Soc. Biol., Paris, 83, 1920, 931.) From acute articular rheumatism. Micrococcus serratus ^Migula. (Xo. 15, Lembke, Arch. f. Hyg., 26, 1896, 309; Migula, Syst. d. Bakt., 2, 1900, 200.) From feces. Winslow and Winslow {loc. cit., 205) regard this as a synonym of Micrococcus candidus Cohn or of Gaffkya tetragena Trevisan. Micrococcus sialosepticus Trevisan. (Coccwssa^u'ariMS sep 1935, 499.) Spheres: Gram-positive. Blood agar: Colonies 0.8 to 1.3 mm, moist and transparent, with crenated edges. Incomplete hemolysis and no alpha-prime appearance. Acid from glucose, lactose and salicin; may foi'm acid from trehalose (doubtful ) . No acid from mannitol or sorbitol. Does not hydrolyze sodium hippurate. Chemical tolerance : Final pH in glu- cose broth 5.1 to 5.4. Does not grow on 10 per cent and 40 per cent bile-blood agar. Action on blood : Incomplete hemol- ysis ; does not produce soluble hemolysin. Doubtful if truly hemolytic strepto- coccus. Fibrinolysin : Does not dissolve human fibrin. Serology : Group K. Aerobe, facultative anaerobe. Source : Human throat. Habitat: Human throat. No indica- tion of relation to disease. 6. Streptococcus acidominimus Ayers and Mudge. (Ayers and Mudge, Jour. Inf. Dis., 31, 1922, 40; S3, 1923, 155.) From M. L., derived to mean a minimum amount of acid. Description taken from Smith and Sherman, Jour. Inf. Dis., 65, 1939, 301. Spheres: Generally occur in short chains. Gram-positive. Gelatin stab : Filiform, beaded growth. No liquefaction. Plain nutrient agar : Small round white colonies. Acid from glucose, lactose and sucrose. May form acid from maltose, trehalose, and mannitol. Sorbitol and salicin usually are not fermented. No acid from arabinose, xylose, raffinose, inulin and glycerol. Sodium hippurate is hydrolyzed but not starch. Carbon dioxide is produced from a 4 per cent peptone-infusion broth. Litmus milk: Little or no visible change. Ammonia is not produced from peptone. Temperature relations : No growth at 10°C. A few cultures grow at 45°C. Do not survive 60°C for 30 minutes. Chemical tolerance : No growth in .01 per cent methylene blue. Growth in 2 per cent but not in 6.5 per cent NaCl. Final pH in glucose broth 6.5 to 5.6. No growth at pH 9.6. Action on blood: No hemolysis, slight greening (alpha). Serology: Negative reaction with serums representing Lancefield groups A, B,C,D,E,FandG. Facultative anaerobe. Distinctive character: Small amount of acidity developed in fermentation tests. Source : Originally 12 cultures were isolated from freshly drawn milk. Also found in bovine vagina, occasionally in the udder, and on the skin of calves. Habitat: Known to occur abundantly in the bovine vagina. The relationship between Streptococcus uberis Diernhofer and other similar strep- tococci is not yet entirely clear. Smith and Sherman (Jour. Inf. Dis., 65, 1939, 301-305) at one time thought that Strep- tococcus uberis and Streptococcus aci- dominimus might be identical. Others have regarded Streptococcus uberis as identical with Group III, Minett (Proc. 12th Internat. Vet. Cong., 2, 1934, 511). Brown (Proc. 3rd Internat. Cong, for Microbiol., 1940, 173) describes a new species, Streptococcus lentus (not Strep- 336 MANUAL OF UETEPari NATIVE T5ACTERIOEOGV iococcus lentus Lehmann, Deutsch. Arch. f. klin. Med., 150, 1926, 144) which belongs to serological group E. He states that a few strains that produced the alpha appearance in blood agar corresponded culturally with Streptococcus nheris. Later Sherman (personal communica- tion) had an opportunity to determine the serological group of several cultures of Streptococcus uberis carefully identi- fied by R. B. Little and found them to belong to Group D. While their char- acters were not exactly the same as the conventional Streptococcus faecalis, he feels that these cultures of Streptococcus uberis were only a variant type of Strep- tococcus faecalis. Appendix II.* The following species of streptococci are listed chiefly because of their historical interest. In many cases the original cultures are lost and their exact taxonomic relationships have not been determined. Bacterium acetylcholini Habs. (Cent, f. Bakt., II Abt., 97, 1937, 194.) From ensilage. Regarded as a stable type of Enter ococcus. Diplococcus homhycis Paillot. (An- nales des Epiphyties, 8, 1922, 131.) From the silkworm (Bombyx mori) . Diplococcus liparis Paillot. (Annales des Epiphyties, 8, 1922, 122.) From larvae of the gJTsy moth (Portheria (Lymantria) dispar). Diplococcus lymantriae Paillot. (Compt. rend. Acad. Sci., Paris, 164, 1917, 526.) From larvae of the gypsy moth (Portheria (Lymantria) dispar). Diplococcus melolonthae Paillot. (Compt. rend. Soc. Biol., Paris, 69, 1917, 57; Annales des Epiphyties, 8, 1922, 118.) From diseased larvae of cock- chafers (Melolontha mclolontha). Diplococcus pieris Paillot. (Annales des Epiphytes, 8, 1922, 128.) From dis- eased caterpillars of the cabbage butter- fly (Pieris brassicae). Diplococcus scarlatinae sanguinis Jamieson and Edington. (Brit. Med. Jour., 1, 1887, 1265.) From the desqua- mation and blood of scarlet fever patients. Enterococcus citreus Stutzer and Wso- row. (Cent. f. Bakt., II Abt., 71, 1927, 117.) From normal pupae of a moth (Euxoa, segetum). Lactococcus agglutinans Plevako and Bakushinskaia. (Microbiology (Rus- sian), 4, 1935,523; abst. in Cent. f. Bakt., II Abt., 9j^, 1936, 64.) Agglutinates baker's yeast. Streptobacillus malae Goadby. (Jour. State Med. London, 30, 1922, 417 ; Strepto- coccus malae Thomson and Thomson, Ann. Pickett-Thomson Res. Lab., 5, 1929, 22.) From the mouth. An aber- rant streptococcus. Streptococcus abortus -equi Hauduroy et al. (Streptococcus abortus equi Ostertag, Monatsh. f. Tierheilk., 12, 1900, 384; Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 508.) From aborting mares. Streptococcus acidi-lactici Chester. (Sphaerococcus acidi lactici Marpmann, Erganzungshefte d. allegemeine Gesund- heitspflege, 2, 1886, 121 ; not Streptococcus acidi lactici Grotenfeldt, Fortschr. d. Med., 7, 1889, 124; Micrococcus lacticus Migula, Syst. d. Bakt., 2, 1900, 66; Ches- ter, Man. Determ. Bact., 1901, 65.) From fresh milk. Streptococcus aerobius Heurlin. (Bakt. Untersuch. d. Keimgehaltes im Genital- kanale der fiebernden Wochnerinnen, Helsingfors, 1910, 60.) From the genital canal . Streptococcus aerogenes Wirth. (Cent. f. Bakt., I Abt., Orig., 95,1926, 290.) From human blood. An aerobic species which produced gas in deep glucose agar. Streptococcus aerophilus Trevisan. (I generi e le specie delle Batteriacee, Milan, 1889, 31 ; not Streptococcus aerophilus Heurlin, Bakt. Untersuch. d. Keimge- haltes im Genitalkanale der fiebernden Wochnerinnen, Helsingfors, 1910, 62.) From air. * Prepared by Miss Eleanore Heist, July, 1938; revised by Prof. Robert S. Breed, New York State Experiment Station. Geneva, New York, February, 1944. FAMILY LACTOBACTERIACEAE 337 Streplococcus alactosus Smith and Brown. (Jour. Med. Res., 31, 1915, 455; Rockefeller Inst, for Med. Res., Mono- graph 9, 1919; Streptococcus haemolyticus II, Holman, Jour. Med. Res., 34, 1916, 377. ) From human tonsillitis ; peritoneal pus. See Manual, 5th ed., 1939, 352 for description of this species. Streptococcus albicans Migula. (Schminkeweisser Streptococcus, Tata- roff, Inaug. Diss., Dorpat, 1891, 69; Mig- ula, Syst. d. Bakt., 2, 1900, 22.) From water. Streptococcus albidus Henrici. (Arb. bakt. Inst. Karlsruhe, 7, Heft 1, 1894,53.) From Cantal cheese. Streptococcus albus Sternberg. (Weis- ser Streptococcus, Maschek, Bakt. Unter- such. d. Leitmeritzer Trinkwasser, Jahr- esber. d. Oberrealschule zu Leitmeritz, 1887; Sternberg, Man. of Bact., 1893, 610; Micrococcus albus Mace, Traite pratique de Bact., 6th ed., 1912, 605.) From water. Streptococcus allantoicus Barker. (Jour. Bact., 40, 1943, 251.) From black mud, San Francisco Baj'. Streptococcus alvearis (Freuss) Trevi- san. {Cryptococcus alvearis Freuss, 1868 ; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 31.) From an infection (foulbrood?) in bees. Streptococcus ambratus Trevisan. (Mi- crococco ambrato, Perroncito and Ajroldi, Giornale d. r. Accad. d. Med. d. Torino, 48, 1885, 809 ; Trevisan, I generi e le specie delle Batteriacee, 1889, 30.) From the respiratory tract of a horse. Streptococcus anhaemolyticus Roily. {Streptococcus anhaemolyticus vulgaris Zangemeister, Munch, med. Wochnschr., 57, 1910, 1268; Roily, Cent. f. Bakt., I Abt., Orig., 61, 1912, 90.) Synonym of Streptococcus saprophyticus Mandelbaum (Ztschr. f. Hyg., 58, 1907, 37; see Brown, Monograph No. 9, Rockefeller Inst. Med. Res., 1919,87). From vaginal secretions, milk and saliva. Streptococcus aphlhicola Trevisan. (I generi e le specie delle Batteriacee, 1889, 30.) From the lesions of foot and mouth disease of cattle. Streptococcus aromaticus van Beynum and Pette. (Directie Landbouw. Vers- lag. Landbouwk. Onderzoek., 42, 1936, 360; also see Hoecker and Hammer, Iowa Agr. Exp. Sta. Res. Bull. 290, 1941, 317.) Produces diacetyl and small amounts of acetylmethylcarbinol in milk. From cream and butter. Streptococcus articulorum Fliigge. (Die Mikroorganismen, 2 Aufl., 1886, 153.) Associated with diphtheria. Trevisan (I generi e le specie delle Bat- teriacee, 1889, 30) considers this identical with Streptococcus diphteriticus Cohn (Beitr. z. Biol. d. Pfianz., /, Heft 2, 1872, 162). Streptococcus asalignus Frost, Gumm and Thomas. (Jour. Inf. Dis., 40, 1927, 703.) From milk. Streptococcus aurantiacus Killian and Feher. (Ann. Inst. Past., 55, 1935, 619.) From Sahara Desert soil. Streptococcus bombycis Sartirana and Paccanaro. (Cent. f. Bakt., I Abt., Orig., 40, 1906, 331; probably not Strep- tococcus bombycis Zopf, Die Spaltpilze, 2 Aufl., 1884, 52.) From diseased silk worms {Bombyx mori). According to Paillot (Les maladies du ver a sole, Lyon, 1928, 171) this is the same as Streptococ- cus pastorianus Krassilstschik. Streptococcus bonvicini Chester. (Streptococcus della leucaemia, Bonvi- cini, Cent. f. Bakt., lA.hi.,21, 1897,211; Chester, Man. Determ. Bact., 1901, 59.) From a case of leucaemia in a dog. Streptococcus bovinus Trevisan. {Mi- crococcus bovinus Trevisan, Rendiconti Reale Inst. Lombardo di Sci. e Lett., Ser. II, 12, 1879; Trevisan, I generi e le specie delle Batteriacee, 1889, 30; not Micro- coccus bovinus Migula, Syst. d. Bakt., 3, 1900, 67; not Streptococcus bovinus Broad- hurst, Jour. Inf. Dis., 17, 1915, 321; not Streptococcus bovinus Frost and Engel- brecht. The Streptococci, 1940, 56.) From human throat; bovine, equine, feline and canine feces. Streptococcus brevis von Lingelsheim. 338 MANUAL OF DETERMINATIVE BACTERIOLOGY (Ztschr. f. Hyg., 10, 1891, 339 and 354.) Not pathogenic. From various human and animal sources. Streptococcus hrevis non hemolyticus Sachs. (Ztschr. f. Hyg., 63, 1909, 466.) From tonsils, vagina and vulva. Streptococcus brightii DeToni and Trevisan. (Streptococcus bei Morbus Brightii, Mannaberg, Cent. f. klin. Med., 9, 1888, 537; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1057; Streptococcals morbi brightii Migula, Syst. d. Bakt., 2, 1900, 28.) From urine of persons suffering from Bright's disease. Probably identical with Streptococcus pyogenes. Streptococcus buccalis Blake. (Jour. Med. Res., 36, 1917, 124.) From the mouth. Proposed to include both Strep- tococcus 7nitis and Streptococcus sali- varius. Streptococcus butyricus (Fitz) DeToni and Trevisan. (Micrococcus butyricus Fitz, Ber. d. deutsch. chem. Gesellsch., Denaeyer Bact. schizom., p. 35; DeToni and Trevisan, in Saccardo, Sylloge Fun- gorum, 8, 1889, 1064.) From bovine feces. Forms butyric acid from calcium lactate. Streptococcus cadaveris Sternberg. (Man. of Bact., 1893, 611.) From liver of yellow fever cadaver. Slreptococctis caprinus Emoto. (Jour. Japan. Soc. Veter. Sci., 3, 1924, 67.) From the cerebro-spinal fluid of goats. Pathogenic for goats. Streptococcus capsulatus Hauduroy et al. (Streptococcus capsulatus gallinarum Dammann and Manegold, Deutsche tier- arztl. Wochnschr., 13, 1905, 577 and Arch. f. Tierheilk., 33, 1907, 41; Haudu- roy ct al., Diet. d. Bact. Path., 1937, 511 ; not Streptococcus capsulatus Binaghi, Cent. f. Bakt., I Abt., 22, 1897, 273.) Causes a disease of chickens. Streptococcus carneus Migula. (Micro- coccus?, List, Inaug. Diss., Leipzig, 1885, 49; Migula, Syst. d. Bakt., 2, 1900, 36.) From the stomach of a sheep. Probably a micrococcus. Streptococcus carnis Chester. (Diplo- coccus, Harrevelt, Cent. f. Bakt., I Abt., 26, 1899, 121; Chester, Man. Determ. Bact., 1901, 60.) From meat. Streptococcus carrosus Lindner. (50 Jubiliiumsber. Westpreuss. Bot.-Zool. Vereins, Danzig, 1928, 254.) From Mexi- can pulque. Streptococcus casei Burri. (Berichte d. Schweiz. Bot. Gesellsch., 51, 1940, 102.) From Emmenthal cheese. Streptococcus caucasicus Migula. (Streptococcus a, von Freudenreich, Cent. f. Bakt., II Abt., 3, 1897, 87; Migula, Syst. d. Bakt., 2, 1900, 42.) From kefir. Possibly related to Strepto- coccus kefir. Streptococcus cerasinus (Lehmann and Neumann) Migula. (Micrococcus?, List, Inaug. Diss., Leipzig, 1885, 17; Kirsch- roter Micrococcus, Adametz, Mitteil. d. osterreich. Versuchssta. f. Brauerei u. Miilzerei in Wien, Heft 1 , 1888, 33 ; Micro- coccus cerasinus siccus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 34 ; Micrococcus cera- sinus Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 179; Migula, Syst. d. Bakt., 2, 1900, 35; not Micrococcus cerasinus Migula, ibid., 170.) From na- sal secretions in a sheep ; also from water. Streptococcus charrini Trevisan. (Mi- crobe de la septicemie consecutive au charbon, Charrin, Compt. rend. Soc. Biol., Paris, 36, 1884, 526; Streptococcus Charrin, Fliigge, Die Mikroorgauismen, 2 Aufl., 1886, 164; Trevisan, I generi e le specie delle Batteriacee, 1889, 30. ) From the organs of a rabbit having anthrax. Streptococcus cinereus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 64.) From water. Streptococcus citreus (Eisenberg) Mig- ula. (Micrococcus?, List, Inaug. Diss., Leipzig, 1885, 60; Cremefarbiger Micro- coccus, Adametz, Mitteil. d. osterreich. Versuchssta. f. Brauerei u. Malzerei in Wien, Heft 1, 1888, 31; Micrococcus cit- reus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 36 ; Migula, Syst. d. Bakt., 2, 1900, 37 ; not Streptococcus citreus Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 179.) FAMILY LACTOBACTERIACEAE 339 From the pancreas of a sheep (List) ; from water (Adametz). Streptococmis ciirophilus van Beynum and Pette. (Directie Landbouw. Vers- lag. Landbouwk. Onderzoek., 4^, 1936, 360; also see Hoecker and Hammer, Iowa Agr. Exp. Sta. Res. Bull. 290, 1941, 317.) Produces diacetyl, acetylmethylcarbinol and volatile acids from citric acid. From cream and butter. Streptococcus coli Migula. (Strepto- coccus coll brevis Escherich, Die Darm- bakterien des Sauglings und ihre Bezieh- ungen zur Physiologie der Verdauung, 1886, 86; Migula, Syst. d. Bakt., 2, 1900, 33.) From stools in cases of infant diar- rhoea. Streptococcus continuosus Black. (Trans. 111. State Dental Soc, 22, 1886, 189.) From the mouth. Streptococcus coronatus (Fliigge) Trevi- san. {Micrococcus coronatus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 175; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 31.) From the air. Streptococcus cuniculi Bergey et al. (Manual, 1st ed., 1923, 50.) From natu- ral infections of rabbits. Morphologi- cally like Streptococcus pyogenes. Streptococcus cystitidis Migula. (Dip- lococcus ureae pyogenes Rovsing, Die Blaseuentziindungen, ihre Aetiologie, Pathogenese und Behandlung, 1890, 39; Migula, Syst. d. Bakt., 2, 1900, 12.) From a case of cystitis. Streptococcus debilis Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 13. ) From meat extracts. Streptococcus dentium (Trevisan) Trevisan. [Micrococcus dentium Trevi- can, Batt. Ital., 1879, 27; Micrococcus foetidus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 172; Trevisan, I generi e le specie delle Batteriacee, 1889, 30. ) From carious teeth. Grows anaerobically in nutrient agar with the production of gas and strong odor. Streptococcus desidens Trevisan. (Mi- crococcus Jlavus desidens Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 177; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 31 ; Micrococcus desidens Migula, Syst. d. Bakt., 2, 1900, 143.) From the air. Streptococcus diacetilactis Matuszew- ski, Pijanowski and Supinska. (Compt. rend. Inst. Bact. Ecole Cent. Agron., Varsovie, Poland, No. 21, 1936, 1.) From milk. Probably Streptococcus cremoris Orla-Jensen. Streptococcus diacetyl aromaticus. (Quoted from Karnad, Indian Jour. Vet. Sci., 9, 1939, 349.) From milk. Pro- duces an aroma in milk cultures. Streptococcus disparts Glaser. (Jour. Agr. Res., 13, 1918, 515.) From the ali- mentary canal of caterpillars. Streptococcus endocarditicus DeToni and Trevisan. (Mikrokokken, Klebs, Arch. f. exper. Pathol., 9, 1878, 52; DeToni and Trevisan, in Saccardo, Syl- loge Fungorum, 8, 1889, 1057.) From cases of endocai'ditis. Streptococcus enteritidis Chester. (Enteritisstreptokokken, Escherich, Jahrb. f. Kinderheilk., 45, 1899, 161; Chester, Man. Determ. Bact., 1901, 59.) From cases of enteritis. Streptococcus enteritis Chester. (Streptococcus, Hirsh, Cent. f. Bakt., I Abt., 22, 1897, 372 and Libman, ibid., 376 : Chester, Man. Determ. Bact., 1901, 56; Streptococcus enteritis var. libmanii Ches- ter, ibid., 66.) From stools in cases of infant diarrhoea. Streptococcus equarius Frost and En- gelbrecht. (A Revision of the Genus Streptococcus, privately published, 1936, 3 pp. and The Streptococci, 1940, 65.) From a throat culture. Streptococcus equinus Lehmann and Neumann. (Streptokokken die sich zu grossen Konvaluten zu-sammenballen, Behring, Cent. f. Bakt., 12, 1892, 194; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 125.) From equine pneumonia. Relation to Streptococcus equinus Andrewes and Horder not clear. Streptococcus felirius Bergey et al. (Manual, 1st ed., 1923, 50.) From natu- ral infections in cats. Morphologically like Streptococcus pyogenes. 340 MANUAL OF DETERMINATIVE BACTERIOLOGY Streptococcus fermenti (Trevisan) Trevisan. {Micrococcus fermenti Trevi- san, Batt. Ital., 1879, 19; Micrococcus viscosus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 172; Trevisan, I generi e le specie delle Batteriacee, 1889, 31.") From a slimy growth in wines. Streptococcus fischeli Chester. (Mi- croorganismvis No. 2, Fischel, Ztschr. f. Heilkunde, 13, 1891, 7 and Cent. f. Bakt., 9, 1891, 611; Chester, Man. Determ. Bact., 1901, 59.) From the blood of in- fluenza patients. Streptococcus foetidus Migula. (Stink- coccus, Klamann, Allegem. med. Central- zeitung, 1887, 1347 ; Diplococcus fluores- cens foetidus Eisenberg, Bakt. Diag., 1891, 10; Migula, Syst. d. Bakt., 2, 1900, 38; Streptococcus fluorescens Chester, Man. Determ. Bact., 1901, 70; Strepto- coccus fluorescens foetidus Miquel and Cambier, Traite de Bact., Paris, 1902, 792.) From cases of ozena. Streptococcus galleriae Chorine. (Compt. rend. Soc. Biol., Paris, 95, 1926, 201.) From the bee moth {Galleria mellonella) . Streptococcus genitalium Dimock and Edwards. (Kentucky Agr. Exp. Sta. Res. Bull. 286, 1928, 162.) Found com- monly in the genital tract of mares. Streptococcus giganteus Migula. (Streptococcus giganteus urethrae Lust- garten and Mannaberg, Vierteljahrsschr. f. Dermatologie u. Syphilis, 1887, 918; Migula, Syst. d. Bakt., 2, 1900, 39.) From human urethra and from urine. Streptococcus gingivae. (Quoted from Annals Pickett-Thomson Res. Lab., 2, 1927, 154.) From human gums and teeth. Streptococcus granulatus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894,55.) From cream cheese. Streptococcus haematosaprus Trevisan. (Mikrokokken der Faulniss, Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 173; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 31.) From putrefying blood. Streptococcus halitus Heim and Schlirf . (Cent. f. Bakt., I Abt., Orig., 100, 1926, 39.) From deposit on the tongue. Streptococcus havaniensis Sternberg. (Man. of Bact., 1893, 612.) From acid vomit of a yellow-fever patient. Streptococcus hemolyticus I, //and /// Holman. (Jour. Med. Res., 34, 1916, 388.) From various human infections. Streptococcus herharum Schieblich. (Cent. f. Bakt., I Abt., Orig., 12^, 1932, 269.) From green plant material. Mo- tile. Related to Streptococcus lactis except it is flagellated. Kolbmuller (Cent. f. Bakt., I Abt., Orig., 133, 1935, 310) identifies this with Enterococcus. Streptococcus hydrophoborum Trevisan. (Streptococcus bei Rabies, Babes, Ztschr. f . Hyg. , 5, 1888, 184 ; Trevisan, I generi e le specie delle Batteriacee, 1889, 30. ) From the brain in a case of rabies. Streptococcus influentiae Trevisan. (I generi e le specie delle Batteriacee, 1889, 30.) From equine influenza. Streptococcus influenzae Thomson and Thomson. (Grippestreptokokkus, Selig- mann, Cent. f. Bakt., I Abt., Ref., 50, 1911, 81 ; Thomson and Thomson, Mono- graph No. 16, Part I, Annals Pickett- Thomson Res. Lab., 1933.) Associated with influenza. Streptococcus infrequens Holman. (Jour. Med. Res., 34, 1916, 388.) From . various human infections. Streptococcus kirchneri Chester. (Dip- lococcus, Kirchner, Ztschr. f. Hyg., 9, 1890, 528; Chester, Man. Determ. Bact., 1901, 57.) From sputum in cases of influenza. Streptococcus kochii Trevisan. (I gen- eri e le specie delle Batteriacee, 1889, 30.) From rabbit septicemia. Streptococcus lacteus Schroter. (Kryp- togam. Flora v. Schlesien, 3, 1, 1886, 149.) From the air and dust. Streptococcus lactis aromaticus Joshi andRam Ayyar. (Indian Jour. Vet. Sci., 6, 1936, 141.) Possibly Streptococcus cremoris Orla-Jensen. From cream. Streptococcus lactis innocuus Stolting. (Inaug. Diss., Kiel, 1935, 16.) From ripening cheese. FAMILY LACTOBACTERIACEAE 341 Streptococcus lagerheimii var. subter- raneum Migula. (Hansgirg, Oesterr. Zeitung, 1888, No. 7 and 8; Migula, Syst. d. Bakt., 2, 1900, 41.) From the wall of a wine cellar. Streptococcus {Diplococcus) lanceolatus avium Gaertner. (Cent. f. Bakt., I Abt., Orig., 54, 1910, 546.) From mastitis in sheep. Streptococcus lapillus Heimand Schlirf. (Cent. f. Bakt., I Abt., Orig., 100, 1926, 39.) From the oral cavity. Streptococcus lentus Lehmann. (Leh- mann, Deutsch. Arch. f. klin. Med., 150, 1926, 144; Streptococcus pyogenes lentus, ibid., 141 ; not Streptococcus lentus Brown, Rept. Proc. Third Internat. Congr. for Microbiol . , Xew York, 1940, 173. ) From urine, cervix, sputum and carious teeth. Streptococcus libaviensis Flatzek. (Cent. f. Bakt., I Abt., Orig., 82, 1919, 240; Bacterium libaviense Flatzek, idem.) From human feces. Motile. Streptococcus lucae Trevisan. (Micro- coccus ulceris mollis de Luca, 1886 ; Trevi- san, Igeneri e le specie delle Batteriacee, 1889, 30.) From chancroidal ulcers. Streptococcus luteus Killian and Feher. (Ann. Inst. Past., 55, 1935, 619.) From Sahara Desert soil. Streptococcus viagnus Henrici. (Arb. bakt. Inst. Ivarlsruhe, i. Heft 1, 1894, 54.) From Brie cheese. Streptococcus malaperti Trevisan. (Mi- crococcus E, Malapert-Neuville, 1887; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 30.) From mineral water of hot springs at Schlangenbad. Streptococcus malignus Trevisan. (Streptococcus pyogenes malignus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 153; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 30.) From a diseased spleen. Probably identical with Strepto- coccus pyogenes. Streptococcus mammitis bovis Hutch- ens. (Hutchens, in Besson, Pract. Bact. Microbiol, and Serum Therapy. Trans, of 5th ed., 1913, 613.) From mastitis in cattle. Streptococcus inargaritaceus Schroter. (In Cohu, Kryptog. Flora v. Schlesieu, 3, 1 , 1886, 149. ) From putrefying blood. Streptococcus mathersi ^NIuslow. (Green producing streptococcus, Tun- nicliff, Jour. Amer. Med. Assoc, 71, 1918, 1733; Mather's coccus, Jordan, Jour. Inf. Dis., 25, 1919, 30; Muslow, ibid., 31, 1922, 295.) From sputum in cases of influenza and pneumonia. Streptococcus maximus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 180.) From a bean and carrot infusion. Streptococcus melanogenes Schlegel. (Berl. tierarztl. Wochnschr., 1906, No. 25, 464.) Produces grayish-yellow pigment in gelatin. Associated with a disease of horses . Streptococcus meningitidis Bonome. (Cent. f. Bakt., 8, 1890, 172 and 703.) From exudates from cases of cerebro- spinal meningitis. Streptococcus merdarius Trevisan. (Streptococcus des selles, Cornil and Babes, Bacteriea, 2nd ed., 1886, 118; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 31.) From feces. Streptococcus microapoikia Cooper, Keller and Johnson. (Amer. Jour. Dis. of Children, 47, 1934, 388 and 596; these authors also use the trinomial Strepto- coccus micro-apoikia enteritis.) From human throat and feces in enteritis in children. See Manual, 5th ed., 1939, .351 for description of this species. Streptococcus rnirabilis Roscoe and Lunt. (Phil. Trans. Roy. Soc, London, 182, 1892, &48.) Streptococcus mixtus Bergey et al. (Manual, 1st ed., 1923, 49.) From a variety of pyogenic inflammations. Streptococcus morbilli Ferry and Fisher. (Jour. Amer. Med. Assoc, 86, 1926, 933.) From blood of persons in early stages of measles. Streptococcus m,orbillosus Trevisan. (Micrococcus morbillosus Trevisan, Ren- diconti Reale Inst. Lombard© di Sci. e Lett., Ser. II, 12, 1879 ; Trevisan, Igeneri e le specie delle Batteriacee, 1889, 30.) From human, canine and porcine measles. Streptococcus murisepticus v. Lingels- 342 MAXUAL OF DETERMIXATIVP: BACTERIOLOGY heim. (Ztschr. f. Hyg., 10, 1891, 331 and 12, 1892, 308.) Migula (Syst. d. Bakt., 2, 1900, 6) considers this a syno- nym of Streptococcus pyogenes. Streptococcus mutans Clarke. (Brit. Jour. Exp. Path., 5, 1924, 142.) Shows extreme variability in colony formation. Regarded as the cause of dental caries. Streptococcus nasalis (Hack) Migula. (Micrococcus nasalis Hack, according to Eisenberg, Bakt. Diag., 3 Aufl., 1891, 55; Migula, Syst. d. Bakt., 2, 1900, 45; Pla7w- coccus yiasalis Migula, loc. cit., 274.) From nasal secretions. Considered motile. Streptococcus necroseos Schroter. (Mi- krokokkus der progressiven Gewebene- krose bei Mausen, Koch, Untersuch. iiber die Atiologie der Wundinfektionskrank- heiten. Leipzig, 1878 or Gesamm. Werke V. Robert Koch, 1, 1912, 87; Schroter, in Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 150; Streptococcus nccroticus Trevi- can, Atti della Accad. Fisio-Medico- Statistica in Milano, Ser. 4, 3, 1885.) From gangrene in mice. Streptococcus nomae Trevisan. (I gen- eri e le specie delle Batteriacee, 1889, 30.) From gangrene of the mouth. Streptococcus non-hemohjticus I, II and /// Holman. (Jour. Med. Res., 5^, 1916, 388.) From various human and animal infections. Streptococcus odontolyticus Belding and Belding. (Dental Items of Interest, 62, 1940, 308.) From dental caries. Later stated by authors (Jour. Amer. Dent. Assoc, 30, 1943, 713) to be a mu- coid variant of Streptococcus salivarius. Streptococcus opacus Heim and Schlirf. (Cent. f. Bakt., I Abt., Orig., 100, 1926, 40.) From the oral cavity. Streptococcus opportunus Brown. (Rept. Proc. Third Internat. Congr. for Microbiol., New York, 1940, 173.) Source not recorded. Belongs to Lance - field's Group B (Sherman, Chase and Niven, Jour. Bact., U, 1941, 101). Streptococcus ovis Weimann. (Ztschr. f. Infectionskrankh. d. Haustiere, 9, 1911, 255.) From infected sheep. Streptococcus pollens Henrici. (Arb. bakt. Inst. Karlsruhe, i, Heft 1, 1894, 57.) From Gouda cheese. Streptococcus pallidus Henrici. (Arb. bakt. Inst. Karlsruhe, i, Heft 1, 1894, 58.) From Neufchatel cheese. Streptococcus parvulus Levinthal. (Cent. f. Bakt., I Abt., Grig., 106, 1928, 195.) From the mucous membrane of the human throat. Streptococcus pastorianus Krassilst- schik. (Compt. rend. Acad. Sci. Paris, 123, 1896, 427.) From silkworms. Streptococcus perniciosus Zopf . (Zopf , Die Spaltpilze, 3 Aufl., 1885, 53; Strepto- coccus perniciosus psittacorum Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 164; Micrococcus perniciosxis Crookshank, Man. of Bact., 3rd ed., 1890, 2b2; Strepto- coccus psi^/acorzrw Migula, Syst. d. Bakt., 2, 1900, 42.) From an infection of par- rots. Found in nodules on the surface of the kidnej's, lungs and spleen. Streptococcus phytophthorus (Frank) Chester. (Micrococcus phytophthorus Frank, Cent. f. Bakt., II Abt., 5, 1899, 134; Chester, Man. Determ. Bact., 1901, 67.) Associated with blight and rot of potato. Streptococcus piima Mac3^ (Jour. Dairy Sci., 6, 1923, 2.) From ropy milk, the Finnish piima. Streptococcus pitycampae a. Dufrenoy. (Compt. rend. Soc. Biol., Paris, 71, 1919, 288.) From processionary moth larvae (Cnethocampa pityocampa). Motile. Gram-positive. Streptococcus pityocampae /3 Dufrenoy (loc. cit.). From processionary moth larvae. Gram-negative. Streptococcus pleomorphus von Wies- ner. (Wien. klin. Wochnschr., 1917, 933 and 1918, 1101; see Lehmann and Neu- mann, Bakt. Diag., 7 Aufl., 2., 1927, 224.) Occurred frequentlj' during the influenza epidemic of 1918. Streptococcus pneumosimilis Frost and Engelbrecht. (The Streptococci, 1940, 57.) From milk and from the throats of dairy employees. Not found in bovine feces. FAMILY LACTOBACTERIACEAE 343 Streptococcus pohjmorphtcs Heim. (Streptococcus, Kraskowska and Nitsch, Cent. f. Bakt., I Abt., Orig., 82, 1918, 264; Heim, see Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 224.) From the throat. Streptococcus productus Prevot. (Compt. rend. See. Biol., Paris, 135, 1941, 105.) An anaerobic streptococcus from a gangrenous lung. Streptococcus proteiformis var. lique- faciens Hauduroy et al. {Enterococcus proteiformis liquefaciens Hauduroy, These Doctorat Med., Strasbourg, 1921 ; Hauduroy et al.. Diet. d. Bact. Path., 1937,521.) From feces. Streptococcus proteus Chester. (Strep- tococcus No. 52, Conn, Report Storrs Agr. Exp. Sta., 1894, 81; Chester, Man. De- term. Bact., 1901, 67.) From cream. Streptococcus pseudohaemolyticus Gum- ming. (Jour. Path, and Bact., 30, 1927, 279.) From sputum of patients with pulmonary tuberculosis. Streptococcus putrefaciens Trevisan. (I generi e le specie delle Batteriacee, 1889, 31 . ) From putrefying blood. Streptococcus pyogenes bovis Lucet. (Ann. Inst. Past., 7, 1893, 325; also see Crookshank, Textbook of Bact., 4th ed., Philadelphia, 1900, 188.) From bovine pus. Streptococcus pyogenes hominis Crook- shank. (Textbook of Bact., 4th ed., Philadelphia, 1900, 187.) From human sources. Streptococcus pyogenes nonhaemolyticus Thomson and Thomson. (Streptocoque pyogen^ nonhemolytique, Weissenbach, Compt. rend. Soc. Biol., Paris, 81, 1918, 819; Thomson and Thomson, Ann. Pick- ett-Thomson Res. Lab., 3, 1927, 183.) From feces and various human infections. Streptococcus radiatus Klein. (Cent, f. Bakt., I Abt., 28, 1900, 417.) From exudate from the udder of a cow. Streptococcus rheumaticus Poynton and Paine. (Poynton and Paine, Lancet, 2, September 22 and 29, 1900, 861 ; Diplo- coccus rheumaticus and Micrococcus rheu- maticus Beaton and Walker, Brit. Med. Jour., January 31, 1903, 237; abst. in Cent. f. Bakt., I Abt., Ref., 55, 1903, 528.) From cases of -rheumatism and endo- carditis. Streptococcus rindfleischii Trevi.san. (Streptococcus bei Mycosis fungoides, Rindfleisch, Deutsche med. Wchnschr., 11 , 1885, 233 ; Trevisan, I generi e le specie delle Batteriacee, 1889, 30.) From skin infection (mycosis fungoides). Streptococcus ruber Lundstrom. (Fin- ska Lakaresallskapets Handlingar, 35, 1893.) Red colonies. Streptococcus rubiginosus Jamieson and Edington. (Brit. Med. Jour., 1, 1887, 1265.) Associated with cases of scarlet fever. Probably identical with Micro- coccus pyogenes Klein, according to Mace (Traite Pratique de Bact., Paris, 4th ed., 1901, 425). Streptococcus rugosus Migula. (Strep- tococcus ureae (non pyogenes) rugosus Rovsing, Die Blasenentziindungen, ihre Aetiologie, Pathogenese und Behandlung, 1890, 44; :Migula, Syst. d. Bakt., 2, 1900, 30; Streptococcus I'ugosus ureae ]\Iiquel and Cambier, Traite de Bact., Paris, 1902, 829.) From cases of cystitis. Streptococcus salivarius brevis and Streptococcus salivarius tenuis Veillon. (Quoted from Thomson and Thomson, Ann. Pickett-Thomson Res. Lab., 3, 1927, 187 and 240.) From the mouth. See Streptococcus tenuis. Streptococcus sanguineus Migula. (Diplococcus pyogenes Pasquale, Giorn. med. d. R. esercito e d. R. marina, 1890; Migula, Syst. d. Bakt., 2, 1900, 36.) From a case of bone tuberculosis. Streptococcus sanguinis Chester. (Streptococcus sanguinis cams Pitfield, Queen's Microscopic Bulletin, Philadel- phia, 1897, 44; Chester, Man. Determ. Bact., 1901, 64.) From the blood of dogs. Streptococcus sanguis White. (Strep- tococcus s.b.e. Loewe, Plummer, Xiven and Sherman, Jour. Amer. Med. Assoc, 130, 1946, 257; White, Thesis, Cornell Cniv., 1946 quoted from White and Xiven, Jour. Bact., 51, 1946, 721.) From 344 MANUAL OF DETERMINATIVE BACTERIOLOGY blood in cases of subacute bacterial endocarditis. Streptococcus saprogenes Trevisan. (I generi e le specie delle Batteriacee, 1889, 31 . ) From putrefying blood. Streptococcus saprophrjticus Mandel- baum. (Ztschr. f. Hyg., 58, 1908, 37.) See Streptococcus anhaemolyticus vul- garis. From mucous membranes. Streptococcus schmidti Trevisan. (Coccus bei Fadenziehende Milch, Schmidt-Mulheim, Arch. f. d. ges. Phys- iol., 27, 1882, 490; Trevisan, I generi e le specie delie Batteriacee, 1889, 31.) From ropy milk. Streptococcus seiferti DeToni and Tre- visan. (Micrococcus bei Influenza, Sei- fert, in Volkmann, Sammlung Klin. Vortrage, 240; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1056.) From sputum and nasal secre- tions of influenza patients. Streptococcus septicus Migula. {Strep- tococcus septicus liquefians Babes, Bakt. Unter. ii. septische Prozesse des Kindes- alters, Leipzig, 1889, 22; Streptococcus septicus liquefaciens Babes, according to Eisenberg, Bakt. Diag., 3 Aufl., 1891, 312 ; Migula, Syst. d. Bakt., 2, 1900, 27; not Streptococcus septicus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 154.) From the blood and organs of a diseased child. Streptococcus septopijaemicus Biondi. (Ztschr. f. Hyg., 2, 1887, 194 and 225.) According to Migula (Syst. d. Bakt., 2, 1900, 6) this is a synonym of Streptococcus pyogenes. From human saliva. Streptococcus sornthalii (Adametz) Migula. (Micrococcus sornthalii Adam- etz, Cent. f. Bakt., II Abt., 1, 1895, 465; Migula, Syst. d. Bakt., 2, 1900, 20.) From milk and hard cheese. Streptococcus sphagni Migula. (Syst. d. Bakt., 2, 1900, 40.) From sphagnum in the Black Forest. Streptococcus sputiyenus Migulu. (Syst. d. Bakt., 2, 1900, 24.) From sputum. Streptococcus stenos Bergey et al. (Manual, 1st ed., 1923, 50.) From a vari- ety of human inflammatory conditions. Streptococcals stramineus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 59.) From Schlosskase. Streptococcus subacidus Holman. (Jour. Med. Res., 84, 1916, 388.) From various human infections. Streptococcus suspectus Trevisan. (Streptococco dell' ematuria, 'Piscia- sangue' dei bovini ; Trevisan, I generi e le specie delle Batteriacee, 1889, 30.) From blood and spleen in cases of bovine hem- aturia. Streptococcus tenuis Veillon. (Arch. Med. Exp. et Anat., 6, 1894, 161.) From human mouth. Streptococcus ierricola van Steenberge. (van Steenberge, Ann. Inst. Past., 34, 1920, 806 ; not Streptococcus terricola Kil- lian and Feher, Ann. Inst. Past., 55, 1935, 619.) From garden soil. Streptococcus toxicatus (Burrill) De- Toni and Trevisan. (Micrococcus toxica- tus Burrill, The Bacteria. Illinois In- dustrial Univ., 11th Ann. Rept., 1882, 42; DeToni and Trevisan, in Saccardo, Syl- loge Fungorum, 8, 1889, 1065.) From diseased plant tissue. Streptococcus trifoliatus Migula. (Diplococcus ureae (non pyogenes) tri- foliatus Rovsing, Die Blasenentziindun- gen, ihre Aetiologie, Pathogenese und Behanglung, 1890, 43; Migula, Syst. d. Bakt., 2, 1900, 29.) From cases of cystitis. Streptococcus turbidus Lehmann and Neumann. (Bouillon tri'ibende Strepto- kokken, Behring, Cent. f. Bakt., 12, 1892, 193; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896,125.) From various human infections, especially erysipelas. Presumably a smooth culture of Strepto- coccus pyogenes. Streptococcus tyrogenus Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 50.) From cheeses. Streptococcus ureae Migula. (Strepto- coccus pyogenes ureae Rovsing, Die Bias- FAMILY LACTOBACTERIACEAE 345 enentziindungen, ihre Aetiologie, Patho- genese und Behandlung, 1890, 45; Migula, Syst. d. Bakt., 2, 1900, 28; not Strepto- coccus ureae Trevisan, I generi e le specie delle Batteriacee, 1889, 31.) From cases of cystitis. Streptococcus urinae Migula. {Diplo- coccus ureae {non pyogenes) Rovsing, loc. cit., 45; Migula, Syst. d. Bakt., 2, 1900, 13.) From cases of cystitis. Streptococcus vaccinae (Cohn) Zopf. (Microsphaeren der Vaccine, Cohn, Arch, f. path. Anat., 55, 1872, 237; Micro- sphaera vaccinae Cohn quoted from Cohn, Beitrage z. Biol, d. Pfianzen, 1, Heft, 2, 1872, 161; Micrococcus vaccinae Cohn, idem; Zopf, Die Spaltpilze, 3 Aufl., 1885, 52.) From lymph of cow po.x pustules. Streptococcus varicellae Trevisan. (Microbio della varicella, Bareggi, 1885, probably in Gazz. Med. ital. Lomb. Mi- lano, 229-242; Trevisan, I generi e le specie delle Batteriacee, 1889, 30. ) From chicken-pox pustules. Streptococcus variolae Trevisan. (Mi- crosphaeren der Variola, Cohn, Arch. f. path. Anat., 55, 1872, 237; Micrococcus variolae Cohn, 1872, quoted from Trevi- san, I generi e le specie delle Batteriacee, 1889, 30.) From lymph of small pox pustules. Regarded by Cohn (Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 161) as a varietj^ of Micrococcus vaccinae Cohn. Streptococcus variolae-ovinae (Plaut) DeToni and Trevisan. {Micrococcus variolae ovinae Plaut, Das organisirte Contagium der Schafpocken und die Miti- gation desselben, Leipzig, 1882; DeToni and Trevisan, in Saccardo, Sylloge Fun- gorum, 8, 1889, 1058.) From the lymph in sheep-pox pustules. Streptococcus vermijormis Sternberg. (Wurmformiger Streptococcus, Maschek, Bakt. Unters. d. Leitmeritzer Trink- wasser, Jahresb. d. Oberrealschule zu Leitmeritz, 1887; Sternberg, Man. of Bact., 1893, 611.) From water. Streptococcus vcrsatilis Broadhurst. (Jour. Inf. Dis., 17, 1915, 323.) From throat of dogs, horse and cattle feces, etc. Streptococcus vini Migula. {Micrococ- cus saprogenes vini II, Kramer, Land- wirtsch. Versuchsstat., 37, 1890, 325 and Die Bakt. in ihren Beziehungen z. Land- wirtsch. u. d. landwirtsch.-technisch. Gewerben, 2, 1892, 140; Migula, Syst. d. Bakt., 2, 1900, 33.) From wine. Streptococcus viscosus Lehmann and Neumann. (Schleimiger Streptokokken, Behring, Cent. f. Bakt., 12, 1892, 193; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 125.) From various human infections. Presumably a mucoid cul- ture of Streptococcus pyogenes. Streptococcus vitulorum Trevisan. (Micrococco della diarrea bianca dei vitel- lini, Perroncito, 1886; Trevisan, I generi e le specie delle Batteriacee, 1889, 30.) From white diarrhoea of calves. Streptococcxis vulgaris Loening. (Munch, med. Wochnschr., 57, 1910, 173 and 247 ; Streptococcus pyogenes vulgaris Thomson and Thomson, Ann. Pickett- Thomson Res. Lab., 3, 1927, 189.) Names applied to Streptococcus pyogenes. Streptococcus weissii Trevisan. (Atti d. Accad. Fisio-Medico-Statistica in Mi- lano, Ser. IV, 3, 1885, 119.) From lung exudate in pleuropneumonia of cattle. Streptococcus zythi Trevisan. (Toru- lacee de la biere malade, Pasteur; Trevi- san, I generi e le specie delle Batteriacee, 1889,31.) From spoiled beer. Streptosiaphylococcus parvulus Heurlin. (Heurlin, Bakt. Unters. d. Keimgehaltes im Genitalkanale der fiebernden Woch- nerinnen. Helsingsfors, 1910, 138.) From genital canal. Intermediate be- tween Streptococcus anaerobius Kronig and Staphylococcus parvulus Veillon and Zuber. 346 Manual of determinative bacteriology Genua 111. Leuconostoc Van Tieghein emend. Hucker and Pedcrson.* (Van Tieghem, Ann. Sci. Nat., 6, Ser. 7, 1878, 170; Belacoccus Orla-Jensen, The Lactic Acid Bacteria. Mem. Acad. Sci. Danemark, Sec. d. Sci., 5, Ser. 8, 1919, 146; Hucker and Pederson, New York Agr. Exp. Sta. Tech. Bui. 167, 1930, 66.) From Latin leucus, clear, colorless; M. L. Nostoc, a genus of blue-green algae. Cells normally spherical. Under certain conditions, such as in acid fruits and vegetables, the cells may lengthen and become pointed or even elongated into a rod. Certain types grow with a characteristic slime formation in sucrose media. Grow on ordinary culture media, but growth is enhanced by the addition of yeast, tomato or other vegetable extracts. Generally, a limited amount of acid is produced, consisting of lactic and acetic acid ; alcohol is also formed, and about one-fourth of the fermented glucose is changed to CO2. Levo lactic acid is always produced, and sometimes dextro lactic acid also. Milk is rarely curdled. Fructose is reduced to mannitol. Habitat: Milk, plant juices. The type species is Leuconostoc mesenteroides (Cienkowski) Van Tieghem. Key to the species of germs Leuconostoc. I. Acid from sucrose. A. Acid from pentoses. 1. Leuconostoc mesenteroides. B. No acid from pentoses. 2. Leuconostoc dextranicum. II. No acid from sucrose. 3. Leuconostoc citrovorum. 1. Leuconostoc mesenteroides (Cien- Spec. Rept. Food Investigation Board, kowski) Van Tieghem. {Ascococcus mes- London, 1923, 134; Leuconostoc pleofructi enteroides Cienkowski, Arb. d. Naturf. Pederson, N. Y. Agr. Exp. Sta. Tech. Gesellsch. a. d. Univ. a. Charkoff, 1878, Bull. 150 and 151, 1929.) From Greek 12; Van Tieghem, Ann. Sci. Nat., (>, Ser. ntesenterium, mesentery; eidus, form 7, 1878, 170; Leuconostoc indicum Liesen- (like). berg and Zopf, Beitr. z. Physiol, u. Probable synonym: Leuconostoc soyae Morph. niederer Organis., Heft 1, 1892, Rpignky, Bull. Sci., Res. Inst, for 19; Streptococcus mesenterioides Migula, Leguminous Crops, Moscow (Russian), Syst. d. Bakt., 2, 1900,25; Leuconostoc f. ^^^^ ^^^ agglutinans Barendrecht Cent. f. Bakt., ' 'gheres: 0.9 to 1.2 microns in diameter, II Abt., 7, 1901, 627; Leuconostoc alter ^ . . . , 1 . , ' ', , ' TT rr, if^ni 1CA occurrmg m pairs and short or Jong Zettnow, Ztschr. f. Hyg., 57, 1907, 154; % ^u u ■ , ., r/ ,, 7 chains. In sucrose solutions the chains Leuconostoc opalanitza Zettnow, toe. , , , , . , , ^. cit ■ Betacoccus arabinosaceus Orla- ^^^ surrounded by a thick, gelatinous, Jensen, The Lactic Acid Bacteria, 1919, colorless membrane consisting of dextran. 152; Leuconostoc arabinosaceus Hoi- Gram-posit ive. land. Jour. Bact., 5, 1920, 223; Glucose gelatin colonies : Small, white Bacillus pleofructi Savage and Hunwicke, to grayish-white, raised, nodular. * Revised by Prof. G. J. Hucker and Prof. Carl S. Pederson, New York State Ex- periment Station, Geneva, New York, September, 1938 ; further revision, December, 1943. FAMILY LACTOBACTERIACEAE 347 Glucose gelatin stab: Growth along entire stab. No liquefaction. Sucrose broth: Abundant growth with massive formation of slimy material. Potato: No visible growth. Indole not formed. Acid from glucose, fructose, galactose, mannose, .xylose, arabinose, sucrose, and generally from lactose, rafiinose, salicin and mannitol. Rarely acid from dextrin, starch, inulin, sorbitol, rhamnose or glycerol. Nitrites not produced from nitrates. Produces slime from sucrose. Most pronounced in sucrose gelatin stab. Aerobic, facultative. Optimum temperature 21° to 25°C. Distinctive characters : Active slime producer in sucrose solutions. Source: Slime in sugar factory. Habitat : Most active of the genus. Encountered in fermenting vegetable and other plant materials. Frequently iso- lated from slimy sugar solutions. 2. Leuconostoc dextranicum (Bei- jerinck) Hucker and Pederson. {Lacto- coccus dextranicus Beijerinck, Folia Microbiologica, Delft, 1912, 377; Beta- coccus hovis Orla-Jensen, The Lactic Acid Bacteria, Copenhagen, 1919, 152 {Leu- conostoc hovis Holland, Jour. Bact., 5, 1920, 223); Streptococcus paracitrovorus Hammer, Research Bui. 63, Iowa Agr. E.xp. Sta., 1920; Hucker and Pederson, N. Y. Agr. Exp. Sta. Tech. Bull. 167, 1930, 67.) From Latin dexter, right; M. L. dextranum, dextran; M. L. dex- tranicus, related to dextran. Note : The description of Strepto- coccus b, V. Freudenreich (Cent. f. Bakt., II Abt., 3, 1897, 47) renamed Strepto- coccus kefir by Migula (Syst. d. Bakt., 2, 1900, 44) is too indefinite to permit the determination of its exact relationship to the organisms in this genus. It is clear, however, that the Streptococcus kefir of these authors and that of Evans (Jour. Agr. Res., 13, 1918, 235) were very similar to if not identical with Leuco- nostoc dextranicum. Streptococcus dis- tendens Hammer (Iowa State Coll. Jour. Sci., 2, 1927, 5) may also be identi- cal with Leuconostoc dextranicum. Spheres : 0.6 to 1.0 micron in diameter, occurring in pairs and in short chains. Gram-positive. Gelatin stab : Gray filiform growth in stab. Agar colonies: Small, gray, circular, slightly raised, entire. Glucose broth : Slight grayish sedi- ment. Litmus milk: Acid, coagulation. Fre- quently shows slight reduction of litmus in bottom of tube. Potato : No visible growth. Indole not formed. Nitrites not produced from nitrates. Produce slime from sucrose in rapidly growing cultures. Acid from glucose, fructose, galactose, maltose, sucrose, and generally from lactose and mannose. No acid from .xylose, arabinose, glycerol, rhamnose, sorbitol, mannitol, starch, rarely raf- finose, inulin or dextrin. Aerobic, facultative. Optimum temperature of growth 21° to 25°C. Distinctive characters : Produces moderate amount of slime in sucrose solutions. Source: Dairy starters. Habitat: Found both in plant mate- rials and in milk products. 3. Leuconostoc citrovoriun (Hammer) Hucker and Pederson. {Streptococcus citrovorus Hammer, Research Bull. No. 63, Iowa Agr, Exper. Sta., 1920; Hucker and Pederson, N. Y. Agr. Exp. Sta. Tech. Bull. 167, 1930, 67.) From Latin citrus, the citron tree; M. L., lemon or orange, hence citric acid; voro, devour. Spheres: 0.6 to 1.0 micron in diameter, occuring in pairs and chains. Gram- positive. Gelatin stab: Filiform growth in stab. No liquefaction. Agar colonies: Small, gray, entire, slightly raised. 348 MANUAL OF DETERMINATIVE BACTERIOLOGY Agar slant: Small, gray, discrete colonies. Glucose broth: Slight gray sediment. Litmus milk : Slightly acid with partial reduction of litmus. Potato: No visible growth. Indole not formed. Nitrites not produced from nitrates. Grows poorly on ordinary media with- out the addition of yeast extract or other growth accessory substance. Acid from glucose, fructose, galactose and lactose. Generally does not form acid from mannose, sucrose, maltose, xylose, arabinose, rhamnose, raffinose, glycerol, dextrin, inulin, starch, salicin, mannitol or sorbitol. Uses citric acid in milk. Aerobic, facultative. Optimum temperature 20° to 25°C. Distinctive character : Non-slime producer. Source: Dairy products. Habitat: Found in milk and dairy products. Appendix: This includes species that probably belong in this genus. The de- scriptions are too meager to permit draw- ing any definite conclusion regarding their relationship to the three species recognized above. Bacterium laevolacticum Migula. (Bacillus acidi laevolaciici Schardinger, Monatsh. f. Chemie, 11, 1890, 544; Migula, Syst. d. Bakt., 2, 1900, 406; Bacterium acidi laevolactici Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 178.) From well water. Leuconostoc lagerheimii Ludwig. (Ludwig, Lehrb. d. niederen Kryptog., 1892, 29; Streptococcus lagerheimii Migula, Syst. d. Bakt., 2, 1900, 41.) From slimy sugar solutions. Micrococcus gelatinogenus Briiutigam. (Pharmaceutische Centralhalle, 1891, No. 30.) From the air. Forms gum in su- crose media. Micrococcus gummosus Happ. (Inaug. Diss., Basel, published in Berlin, 1893, 31.) From slimy sugar solutions. Myxococcus betae Gonnermann. (Oesterr.- Ungar. Ztschr. f. Zuckerind. u. Landw., 136, 1907, 883.) From sugar beet juice. Streptococcus citrovorus-paracitrovorus Vas and Csiszdr. (Milchwirtsch. Fortsch., 18, 1936, 68.) From cream and butter. Streptococcus hornensis Boekhout. (Cent. f. Bakt., II Abt., 6, 1900, 162.) From slimy, sweetened condensed milk. A strong dextran former. Related to Leuconostoc mesenteroides . Zoogloea termo Cohn. (Cohn, Nov. Act. Acad. Caes. Leop. -Carol. Nat. Cur., 2■l^, 1854, 123.) The only species in the genus Zoogloea as originally proposed. From running water. Scheibler used this name for a zoogloea-forming organism from slimy sugar solutions in Neue Ztschr. f. Riibenzucker- Ind., /, 1878, 366 and prob- bably also in Ztschr. d. Vereins f . Riiben- zucker- Ind., 1874, 330. The latter refer- ence apparently is not available in America. See Buchanan, General Syst. Bact., 1925, 530 for a history of the genus Zoogloea. FAMILY LACTOBACTEKIACEAE 349 TRIBE II. LACTOBACILLEAE WINSLOW ET AL. (Jour. Bact., 5, 1920, 211.) Rods, often long and slender. Non-motile. Gram-positive. Pigment formation rare. When present, yellow or orange, to rust or brick-red in color. Poor surface growth (except in genus Microhaclerium) because these bacteria are generally micro- aerophilic or anaerobic. Carbohydrates and polyalcohols are changed either by homofermentation to lactic acid or by heterofermentation to lactic, acetic, propionic or butyric acids, alcohol and carbon dioxide. Growth on potato is poor or absent. Gelatin is not liquefied. Nitrates are not reduced (except in genus Microhaclerium) . Several species grow at relatively high temperatures. May or may not produce catalase. Key to the genera of tribe Lactobacilleae. I. Always produce lactic acid from carbohydrates, a. Catalase negative. Microaerophilic. Genus I. Lactobacillus, p. .349. aa. Catalase positive. Aerobic. Genus II. Microbacteriuin, p. 370. II. Ferments carbohydrates, polyalcohols and lactic acid with the formation of propionic and acetic acids, and carbon dioxide. Catalase positive. Genus III. Propionibacterium, p. .372. III. Ferments carbohydrates, polyalcohols and lactic acid with the formation of butyric and acetic acids, and carbon dioxide. Generally catalase negative. Genus IV. Bv.tyribacterium,, p. .379. Germs I. Lactobacillus Beijcrinck.* (? Dispora Kern, Biol. Zent., 2, 1882, 135; ?Tyrothrix Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 79; ?Pacinia Trevisan, Atti della Accad. Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 83; ?Saccharobacillus van Laer, Contributions al'histoire des ferments des hydrates de carbone. Mem. Acad. Royale de Belgique, 43, 1889; Lacto- bacter Beijerinck, Cent. f. Bakt., II Abt., 6, 1900, 200; Beijerinok, Arch, neerl. d. sci. exact, et nat. Hadrlem, Ser. 2, 7, 1901, 212; Streptobacillus Rist and Khoury, Ann. Inst. Past., 16, 1902, 70; ?Brachijbacterium Troili-Petersson, Cent. f. Bakt., II Abt., 11, 1903, 138; Caseobaclerium Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, .336; Plo- cajnobacterium Lowi, Wiener klin. Wochnschr., 33, 1920, 7.30 (in part) ; Lactubacterium van Steenberge, Ann. Inst. Past., 34, 1920, 806; Bifidobacterium Orla-Jensen, Le Lait, 4, 1924, 469; Acidobacterium Heim, see Schlirf, Cent. f. Bakt., I Abt., Orig., 97, 1925, 111: Bifidibacterium Prevot, Ann. Inst. Past., 60, 1938, 303.) From M. I.., lactic and bacillus, a rod. Rods, usually long and slender. Microaerophilic Carbohydrates and poly-alco- hols are changed by homofermentalion to lactic acid, or by heterofermentation to lactic and acetic acids, alcohols and carbon dioxide. Catalase negative. Found in fermenting animal (especially dairy) and plant products. The type species is Lactobacillus caucasicus Beijerinck. * Completely revised by Prof. Carl S. Pederson, New York State Experiment Sta- tion, Geneva, New York, in consultation with Prof. .1. M. Sherman, Cornell Uni- versity, Ithaca, New York and Prof. L. F. Rettger, Yale University, New Haven, Conn., June, 1938; further revision by Prof. Carl S. Pederson, January, 1945. 350 MANUAL OF DETERMINATIVE BACTERIOLOGY Key to the species of genus Lactobacillus. I. Produce only traces of by-products other than lactic acid. Homofermentative. A. Optimum temperature 37° to 60°C or higher. Sub-genus Thermobacterium Orla-Jensen (The Lactic Acid Bacteria, 1919, 160). 1. Acid from lactose. a. Optimum temperature 37° to 45°C. b. Produce levo lactic acid. 1. Lactobacillus caucasicus. 2. Lactobacillus lactis. bb. Produce inactive or dextro lactic acid. c. Microaerophilic. 3. Lactobacillus helveticus. 4. Lactobacillus acidophilus. cc. Anaerobic in freshly isolated cultures. 5. Lactobacillus bifidus, aa. Optimum temperature 45° to 62°C; usually no acid from maltose. 6. Lactobacillus bulgaricus. 7. Lactobacillus thermophilus . 2. No acid from lactose. 8. Lactobacillus delbrueckii. ]i. Optimum temperature 28° to 32°C. Sub-genus Streptobacteriuvi Orla-Jensen {loc. cit., 166). 1. Acid from lactose. a. Produces de.xtro lactic acid. Often prefers lactose to sucrose and maltose. 9. Lactobacillus casei. aa. Produces inactive lactic acid. 10. Lactobacillus plantarum. 2. No acid from lactose. 11. Lactobacillus Icichnianyiii . II. Produce considerable amounts of by-products other than lactic acid (carbon dioxide, alcohol and acetic acid ; mannitol from fructose) . Heterofermentative. Sub-genus Betabacterium Orla-Jensen {loc. cit., 175).* A. Optimum temperature 28° to 32°C. Usually ferment arabinose. 1 . Does not ferment rafhnose, and usually does not ferment sucrose or lactose. 12. Lactobacillus brevis. 2. Ferment raffinose, sucrose and lactose. 13. Lactobacillus buchneri. 14. Lactobacillus pastorianus. B. Optimum temperature 35° to 40°C or higher. Usually does not ferment arabinose. 15. Lactobacillus Jermenti. * Also see discussion of Betabacterium caucasicum, p. 358. FAMILY LACTOBACTERIACEAE 351 1. Lactobacilus caucasicus Beijerinok. {Bacillus caucasicus Beijerinck, Arch. n^erl. d. sci. exact, et nat., 23, 1889, 428; Beijerinck, ibid., S^r. 2, 7, 1901, 212; not Bacillus caucasicus v. Freudenreich, Cent. f. Bakt., II Abt., 3, 1897, 54 and 135; Bacterium caucasicus Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 130; not Bacterium caucasicum Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 209; not Betahacterium caucasicum Orla- Jensen, The Lactic Acid Bacteria, 1919, 175.) From Greek Caucasia, M. L. caucasicus, of the Caucasus. The following is a possible or probable synonym : Streptobacillus Ichenis Rist and Khoury, Ann. Inst. Past., / 6, 1902, 70. Description taken from the two reports of Beijerinck {loc. cit.). Rods : Thin and variable in size, occur- ring singly or in filaments. Xon-motile. Non-spore-forming. Gram-positive (not recorded in early descriptions). Gelatin : No liquefaction. Wort gelatin : Small, white colonies. Agar colonies : Small . Broth : Carbohydrates necessary for growth. Milk: Rapid acid production with coagulation, no action in casein. Utilizes animal peptones with diffi- culty, utilizes vegetable peptones more readily. Acid from glucose, sucrose, maltose and lactose. No action on starch. Action on other carbohydrates not studied. Lac- tose in milk converted to levo lactic acid with little carbon dioxide. Microaerophilic. Optimum temperature 40° to 44°C. Temperature range 25° to 45°C. Source : From kefir and cheese. Habitat: Occurs symbiotically with yeast in kefir. Prototype: Dispora caucasica Kern. (Kern, Biol. Zent., 2, 1882, 135; later in Bull, de la Soc. Imp. des Naturalistes de Moscow, 56, 1882, 168; Bacterium cau- casicum Zopf, Die Spaltpilze, 3 Aufl., 1885,90; Bacillus kaukasicus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 270; Faciiita caucasica Trevisan, I generi e le specie delle Batteriacee, 1889, 23.) The description bj^ Kern of an organism from kefir grains is confused probably because the organism (a spore former) which he isolated by the use of Cohn's solution was not the same as the presum- ably granulated Lactobacillus he saw in microscopical preparations of kefir. Beijerinck was apparently the first to have isolated a lactobacillus from kefir in pure culture and to have given a suffi- ciently complete description to make reidentification possible. It should be noted that from the characters given, this could not have been the same species as that isolated later from kefir by v. Freudenreich {loc. cit.) and Orla- Jensen (loc. cit.). 2. Lactobacillus lactis (Orla-Jensen) Holland. (Bacillus Inctis acidi Leich- mann. Cent. f. Bakt., II Abt., 2, 1896, 779 ; Milch. Zeitung, 25, 1896, 67 ; Thcrmo- bacterium lactis Orla-Jensen, The Lactic Acid Bacteria, 1919, 164; Lactobacillus laciis-acidi Holland, Jour. Bact., 5, 1920, 223; Holland, ideiii.) From Latin lac, milk. Henneberg (Handb. der Garungsbakt., 2 Aufl., 2, 1926, 128) regards Bacilhis lactis acidi Leichmann as identical with Thermobacterium lactis Orla-Jensen. Rods : Long forms with a tendency to grow into threads, often strongly curling. Occur singly or in pairs in young vigorous cultures. Generally contain volutin grains. Gram-positive (not recorded in original description). Milk : Acid produced followed by coagulation in one to four days. 1.7 per cent acid produced. Acid from fructose, glucose, mannose, galactose, sucrose, maltose, lactose, raffi- nose and dextrin. Glycerol, xylose, arabinose, rhamnose, sorbitol, mannitol, inulin and starch not fermented. Salicin may or may not be fermented. Forms levo lactic acid with only a trace of other products. Temperature relations : Optimuna 40°C. 352 MANUAL OF DETERMINATIVE BACTERIOLOGY Minimum 18° to 22°C. Maximum 50°C'. Source : From milk and cheese. Habitat : Undoubtedly widely distrib- uted in milk or milk products. 3. Lactobacillus helveticus (Orla-Jen- sen) Holland. {Bacillus e, von Freuden- reich, Cent. f. Bakt., II Abt., 1, 1895, 173; also Landw. Jahrb. d. Schweiz, 1895, 211 ; BacUbis casei e, v. Freudenreich and Thoni, Landw. Jahrb. d. Schweiz, 1904, 526; also Orla-Jensen, Cent. f. Bakt., II Abt., 13, 1904, 609; Caseobacterhmi e, Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 337; Thermobacterivm helveti- cwn Orla-Jensen, Maelkeri-Bakteriologie, 1916, 35; also the Lactic Acid Bacteria, 1919, 164; Bacterium casei e, Holland, Jour. Bact., 5, 1920, 221; Holland, ibid., 223.) From Latin helveticus, Swiss. Rods: 0.7 to 0.9 by 2.0 to 6.0 microns, occurring singly and in chains. Non- motile. Gram-positive. Whey gelatin colonies : Does not grow readily at temperatures required for incubation of gelatin. Lactose agar colonies: Small, grayish, viscid. Milk: Acid, with coagulation; may become slimy. Nitrites not produced from nitrates. Acid from glucose, fructose, galactose, mannose, maltose, lactose, and smaller amounts from dextrin. The lactic acid is inactive. Temperature relations : Optimum 40° to 42°C. Minimum 20° to 22°C. Maxi- mum 50°C. Microaerophilic . Source : From sour milk and cheese. Habitat: Widely distributed in dairy products. 4. Lactobacillus acidophilus (Moro) Holland. {Bacillus acidophilus Moro, Wiener klin. Wochnschr., 13, 1900, 114; also Jahrb. f . Kinderheilkunde, 52, 1900, 38; Holland, Jour. Bact., 5, 1920, 215; Plocamobacterium acidophilum Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 510; Thermobacterium intestinale Orla-Jensen, Orla-Jensen and Winther, Cent. f. Bakt., II Abt., 93, 1936, 321.) From Latin acidus, sour; M. L. acidus, acid and Greek philus, loving. Possible synonyms : Milchsaurebacil- lus, Boas and Oppler, Deutsche med. Wochnschr., 21, 1895, 73; Diagnostik und Therapie d. Magenkrankheiten, II Teil, 1907, 265 {Lactobacillus boas-oppleri Bergeyetal., Manual, 1st ed., 1923,243); Bacilbis exilis Tissier, La flore intestinale des nourrissons, Paris, 1900, 102; Bacillus gastrophilus Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 424 {Bac- terium gastrophilum Lehmann and Neu- mann, Bakt. Diag., 5 Aufl., 2, 1912, 305) ; Bacillus acetogenus a Distaso, Cent. f. Bakt., I Abt., Orig., S9, 1911, i9; Bacillus acetogenus j8 Distaso, ibid., 51; Bacillus acetogenus proteiformis Distaso, ibid., 52; Bacillus acetogenus exilis Distaso, ibid., 53; Bacillus paraexilis Distaso, ibid., 66; Bacillus dimnrphus Distaso, ibid., 55; Bacillus dimorphus var. longa Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 440; {Bacteroides dimorphus Bergey et al., Manual, 1st ed., 1923, 258); Streptobacil- lus long us Distaso, ibid., 439; Thermo- bacterium acidophilum Henneberg, Cent. f. Bakt., II Abt., 91, 1934, 102. Description of Moro supplemented by material from Kulp and Eettger, Jour. Bact., 9. 1924, 357; Curran, Rogers and Whittier, Jour. Bact., 25, 1933, 595; and Rcttger, Levy, Weinstein and Weiss, Lactobacillus acidophilus, Yale Univ. Press, New Haven, 1935. Rods : 0.6 to 0.9 by 1.5 to 6.0 microns, occurring singly, in pairs and in short chains with rounded ends. Non-motile. Dimensions variable (Kulp and Rettger), (Curran, Rogers and Whittier). Gram- positive; old cultures often Gram-nega- tive (Moro). Gelatin : No growth at 20°C. No lique- faction. Wort-agar (Moro ) or tomato agar (Kulp and Rettger) plates: Surface colonies, peripheries a capilliform maze of long, delicate, twisted, fuzzy projections, cen- ter appears as a thick, dark, felt-like FAMILY LACTOBACTERIACEAE 353 mass. Deep colonies, small, irregularly shaped, with fine radiate or ramified projections. Wort-agar slants : Growth scanty, limited, dry, veil-like. Wort -broth : After 48 hours, fine, floccu- lent sediment- Other acid broths sedi- ment whitish, slight turbidity. Milk: Slow growth with small inocu- lum. Coagulates from the bottom up Potato: No growth. Acid but no gas from glucose, sucrose and lactose (Moro). Acid from glucose, fructose, galactose, mannose, maltose, lactose and sucrose. Some cultures ferment raffinose and trehalose and have slight action on dextrin. Xylose, arabi- nose, rhamnose, glycerol, mannitol .sorbitol, dulcitol and inositol not fer- mented (Kulp and Rettgerj. Inactive lactic acid and volatile acids formed from sugars (Curran, Roger.s and Whittier). No visible growth in carbohydrate -free media (Rettger, Levy, Weinstein and Weiss) . Optimum temperature 37°C. No growth at 20° to 22°C (Moro). Maximum temperature 43° to48°C (Curran, Rogers and Whittier). Not pathogenic for laboratory animal.>^. Microaerophilic . Distinctive characters : Grows in acid media. Unless frequent transfers are made, organism may become Gram-nega- tive and rapidly develop characteristic degeneration forms (Moro). The so- called original strains of Bacillu-s om pus of ears of scarlet fever patients. Corynebacterium avidum (Eggerth) Prevot. (Bacteroidcs avidus Eggerth, Jour. Bact., 30, 1935, 289; Prevot, Ann. Inst. Past., 60, 1938, 304.) Forms gas in some media. From the human intes- tine. Pederson (Jour. Bact., 50, 1945, 478) secured a culture of this species from Eggerth, and found that it fer- mented glucose with the production of higher fatty (presumably butyric) acids, and lactic acid. The species should probably be placed in Butyribacterium Barker. Corynebacterium blattellae Glaser. (Jour. Exp. Med., 5/, 1930, 907.) Found in the fat lx)dy of the German cockroach ( Blattella germanica). For a more com- plete description see Manual, 5th ed., 1939, 978. FAMILY CORYNEBACTERIACEAE 403 Corijnebacteriuin bruneum Kisskalt and Berend. {Bacterium hninexim y ar- bor escens, quoted from Kisskalt and Berend, Cent. f. Bakt., I Abt., Orig., 81, 1918, 446 ; Kisskalt and Berend, idem.) Source not given. Corynebacterium cerebralis Eberson. (Jour. Inf. Dis., 23, 1918, 17.) From the brain in a case of meningitis. Corynebacterium. ceruminis (Graham- Smith) Eberson. {Bacillus ceruminis Graham-Smith, Jour. Hyg., 4, 1904, 258; Eberson, Jour. Inf. Dis., 23, 1918, 8.) Indole is not formed. From normal and scarlet fever-infected ears. Corxjnebacterium commune Martin. (Compt. rend. Soc. Biol., Paris, 81, 1918, 991 and 998.) From the pharynx. Corynebacterium cremoides (Lehmann and Neumann) Jensen. {Bacterium cre- moides Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 253; Jensen, Proc. Linn. Soc. New So. Wales, 59, 1934, 40.) From tapwater, Wiirzburg. Lehmann and Neumann recognize this species as a Corynebacterium in the seventh edition of their determinative bacteriologj' (Bakt. Diag., 7 Aufl., 2, 1927, 710) but do not use the binomial Corynebacterium cremoides except in the index, page 848. Jensen has reisolated this organism from soil in Australia. Bacterium cocciforme Migula (Kultur No. 2, Severin. Cent. f. Bakt., II Abt., 1, 1895, 160; Migula, Syst. d. Bakt., 2, 1900, 439) from manure is regarded by Jensen {loc. c.it.) as closely related to this species. Corynebacterium cuculi (Graham- Smith) Bergey et al. {Bacillus cuculi Graham-Smith, Jour, of Hyg., 4, 1904, 315; Bergey et al., Manual, 1st ed., 1923, 387.) From the throat of a cuckoo. For a more complete description see IManual, 5th ed., 1939, 802. Corynebacterium. cuniculi Hauduroy et al. {Bacillus pyogenes cuniculi Com- inotti, Clinica Veterinaria, 44, 1921, 45; Hauduroy et al., Diet. d. Bact. Path., 1937, 147). Reported as Gram-variable by Cominotti, as Gram-negative by Hauduroy et al. Causative agent of suppurative infection of rabbit. Corynebacterium cutis Hauduroy et al. {Bacillus cutis communis Nicolle, quoted from Costa, Troisser and Dau- vaugne, Compt. rend. Soc. Biol., Paris, 81, 1918, 1003; Bacillus cutis Costa, Troisser and Dauvaugne, ibid., 1004; Bacterium cutis commune Nicolle, quoted from Debre and Letulle, La Presse Med., 27, 1919, 515; Hauduroy et al., Diet. d. Bact. Path., 1937, 148). From normal skin and nasal passages. Corynebacterium delicatum Eberson. (Jour. Inf. Dis., 23, 1918, 16.) From ascitic fluid. Also from blood. Corynebacterium dermophilum (Rohde) Andrewes et al. {Bacillus dermophilus Rohde, Miinch. med. Wchnschr., 68, 1921, 234; Andrewes, Bulloch, Douglas, Dreyer, Fildes, Ledingham and Wolf, Diphtheria, London, H. M. Stationery Office, 1923, 391.) From the skin. Corynebacterium diphtheroides Prevot. (Bacille diphtheroide, Jungano, Compt. rend. Soc. Biol., Paris, 61, 1909, 112; Prevot, Ann. Inst. Past., 60, 1938, 304.) Forms gas in some media. From the intestines of white rats. Corynebacterium epidermidis Eberson. (Jour. Inf. Dis., 23, 1918, 17.) From skin and pus pockets. Resembles Cory- nebacierium suppuratum Eberson. Corynebacterium flocculens Eberson. (Jour. Inf. Dis., 23, 1918, 17.) From a case of appendicitis. Corynebacterium gallinarum Bergey et al. {Bacillus diphtheroides galli- narum Graham-Smith, Jour, of Hyg., 4, 1904, 314; Bergey et al.. Manual, 1st ed., 1923,387.) From the throats of chickens. For a more complete description see Manual, 5th ed., 1939, 802. Corynebacterium glandulae Eberson. (Jour. Inf. Dis., 23, 1918, 14.) From lymph glands in Hodgkin's disease but not specific for the disease. Corynebacterium granulomatis maligni de Negri and Mieremet. (Cent. f. Bakt., I Abt., Orig., 68, 1913, 292.) Causative agent of human malignant granuloma. 404 MANUAL OF DETERMINATIVE BACTERIOLOGY Corynebacterium granulosum Prevot. (Bacille granuleux, Jungano, Compt. rend. Soc. Biol., Paris, 66, 1909, 123; Pr6vot, Ann. Inst. Past., 60, 1938, 304.) From the intestines of white rats. Corynebacterium hepatodystrophicans (Kuczinski) Prevot. (Bacillus hepato- dystrophicans Kuczinski, Der Erreger des Gelbfiebers-Wesen und Wirkung, Monographic, 1929, Berlin; Prevot, Ann. Inst. Past., 60, 1938, 304.) Manteufel (Cent. f. Bakt., I Abt., Orig., 138, 1937, 309) regards this species as identical with Bacillus renale (cunicidi) Manteufel and Herzberg. Common in the organs of monkeys infected with yellow fever virus. Corynebacterium hodgkinii Bunting and Yates. (Bunting and Yates, Arch. In- ternal Med., 12, 1913, 236 ; Johns Hopkins Hosp. Bull., 25, 1914, 173; Bacillus hodgkini Mellon, Jour. Bact., 2, 1917, 271; Fusiform is hodgkini Holland, Jour. Bact., 5, 1920, 223.) From lymph glands in Hodgkin's disease. Not pathogenic. Thought by Fox (Jour. Med. Res., 32, 1915, 309) and Eberson (Jour. Inf. Dis., 23, 1918, 11) not to represent a definite species. Eberson recognized four separ- ate species isolated from human lymph glands, three being from glands in Hodg- kin's disease (Corynebacterium aurantia- cum, C. pseudodiphtheriae, C. glandulae and C. lymphophihim) . Corynebacterium liquefaciens Prevot. {Bacillus parvus liquefaciens Jungano, Compt. rend. Soc. Biol., Paris, 65, 1908, 618; Prevot, Ann. Inst. Past., 60, 1938, 304; not Corynebacterium liquefaciens Andrewes et al., Diphtheria, London, 1923, 408; not Corynebacterium lique- faciens Jensen, Proc. Linn. Soc. New So. Wales, 59, 1934, 49.) From human in- testine. Corynebacterium liquefaciens Andrewes et al. (Bacillus diphtheroides liquefa- ciens Graham-Smith, Jour. Hyg., 4, 1904, 258; Bacillus liquefaciens Mellon, Jour. Bact., 2, 1917, 290; Andrewes, Bulloch, Douglas, Dreyer, Fildes, Ledingham, and Wolf, Diphtheria, London, 1923, 408 . ) From mouth of a patient . Motile . Corynebacterium lymphae vaccinalis Levy and Tickler. (Deutsch. med. Wchnschr., 26, 1900, 418; Corynebacte- rium pyogenes Lewandowsky, Cent. f. Bakt., I Abt., Orig., 36, 1904,473.) From animal lymph. Corynebacterium lymphophilum (Tor- rey) Eberson. (Bacillus lymphophilus Torrey, Jour. Med. Res., 34, 1916, 79; Eberson, Jour. Inf. Dis., 23, 1918, 23.) Anaerobic . From lymph glands in Hodg- kin's disease, but not specific for the disease. Corynebacterium maculatum (Graham- Smith) Ford. (Bacillus maculatus Graham-Smith, Jour. Hyg., 4, 1904, 258; Ford, Textb. Bact., 1927, 277.) From throat. Regarded as a Corynebacterium by Eberson (Jour. Inf. Dis., 23, 1918, 7). Corynebacterium metritis Hauduroy et al. (Souckin, Sovetskaia Veter., No. 11, 1934; Hauduroy et al., Diet. d. Bact. Path., 1937, 156.) Causative agent of metritis in rabbit. Corynebacterium miltinum Kisskalt. (Quoted from Kisskalt and Berend, Cent. f. Bakt., I Abt., Orig., 81, 1918, 446). Source not given. Corynebacterium nodosum (Migula) Eberson. (Bacillus nodosus parvus Lustgarten-Mannaberg, Vierteljahr- schrift f. Dermatol, u. Syphilis, 1887, 914; Bacterium nodosum Migula, Syst. d. Bakt., 2, 1900, 416; Eberson, Jour. Inf. Dis., 23, 1918, 4.) Found in the normal human urethra. Corynebacterium nubilum (Frankland and Frankland) Jensen. (Bacillus nu- bilus Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 386; Bacterium nubilum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 255; Chromobacterium nu- bile Ford, Textb. of Bact., 1927, 472; Flavobacterium nubilum, incorrectly as- cribed to Bergey, by Jensen, Proc. Linn. Soc. New So. Wales, 59, 1934, 44; Jensen, idem.) From water and soil. The iden- tity of this species is doubtful. The original description by the Franklands is incomplete. Zimmerman (Bakt. un- serer Trink- u. Nutzwasser, Chemnitz, 1, FAMILY CORYNEBACTERIACEAE 405 1890, 28) thought he found the same or- ganism and described it as Gram-nega- tive. Lehmann and Neumann (Bakt. Diag., 1 Aufl., 2, 1896, 255) who studied one of Zimmermann's cultures reported this culture as Gram-positive and non- motile, while the Franklands and Zim- mermann speak of an active, circular motility of the very slender rods. Leh- mann and Neumann later (Bakt. Diag., 7 Aufl., 2, 1927, 710) list their Bacterium nubilum (with other Gram-positive, non- motile rods) as a possible Corynebacte- rium. Jensen failed to find anything that exactly corresponded to anj' of these descriptions but describes a small, Gram-jx)sitive, jxjorly-growing, pink to red, slow gelatin-liquefying rod which he says has little in common with corj^ne- bacteria as a new variety Corynebacte- rium nubilum var. nanum. Because the earlj' cultures developed rhizoid growths in stiff gelatin before liquefaction, Zim- mermann originally planned to call this species Bacillus nebulosus {loc. cit., 29), a name that has been used by later au- thors for several different organisms. Attention should be called also to Bacil- lus caudalus Wright, an organism which Conn found to show occasional motility (polar) and named Pseudomonas cauda- datus. This common, slender, gelatin- liquefying. Gram-negative, white to yel- low chromogenic rod is much like the Franklands' and Zimmermann's organ- ism (see Conn, Xew York State Exp. Sta. Tech. Bull. 67, 1919, 38). Corynebacterium paralyticans (Robert- son) Ford. {Bacillus paralyticans Robert- son, Rev. Neurol, and Psychiat., Edinburgh, 1, 1903, 470; Ford, Textb. of Bact., 1927, 281.) From cerebro- spinal fluid. A diphtheroid. Thought at one time to be the causal agent of general paralysis. Corynebacterium parvum Prevot. {Corynebacterium parvum infectiosum Mayer, Cent. f. Bakt., I Abt., Grig., 98, 1926, 370; Prevot, Man. de Class, et Determ. des Bact^ries Ana^robies, Mono- graphie, Inst. Past., Paris, 1940, 202.) From blood in a post-natal fever. Corynebacterium periplanetae Bergey et al. {Corynebacterium periplanetae var. americana Glaser, Jour. Exp. Med., 51, 1930, 59; Bergey et al., Manual, 4th ed., 1934, 550.) Found in the fat body of the American cocki'oach {Periplaneta americana). For a more complete de- scription see Manual, 5th ed., 1939, 798. Corynebacterium plumosum (Fox) Ford. {Mycobacterium- plumosum Fox, Cent. f. Bakt., I Abt., Grig., 70, 1913, 148; Ford, Textb. Bact., 1927, 281.) From blood of patient with chronic endo- carditis. Corynebacterium pse udodiphtheriae Eberson. (Jour. Inf. Dis., 23, 1918, 14.) Hemoglobinophilic. From tonsils. Corynebacterium putidum Eberson. ( Bacillus diphtheroides liquefaciens Graham-Smith, Jour. Ilyg., 4, 1904, 258; Eberson, Jour. Inf. Dis., 23, 1918, 16.) From mouth. Cultures described bj^ Graham-Smith liquefied gelatin and were sluggishly motile. Corynebacterium pyogenes bovis (Roux) Prevot. {Bacillus pyogenes bovis Roux, Cent, f . Bakt., I Abt., Orig., 34, 1905, 541 ; Eubacterium pyogenes bovis Prevot, Ann. Inst. Past., 60, 1938, 295; Prevot, Man. de Class, et Determ. des Bacteries Anaer- obies, Monographie, Inst. Past., Paris, 1940, 204.) Common in bovine suppura- tions. Said by Roux to be identical with Bacillus pyogenes bovis Kunnemaun. Prevot says it is probably identical with the pyogenic Corynebacterium of Lucet. See Corynebacterium pyogenes Eberson. Corynebacterium rcnale cuniculi Prevot. {Bacterium renale and Bacte- rium renale {cuniculi) Manteufel and Herzberg, Cent. f. Bakt., I Abt., Grig., 116, 1930, 266; Bacillus renale and Bacillus renale {cuniculi) Manteufel, ibid., 138, 1937, 306; Prevot, Ann. Inst. Past., 60, 1938, 304.) Gram- variable. Forms gas in some media. From rabbit kidneys. Corynebacterium ruedigeri (Mellon) 406 MANUAL OF DETERMINATIVE BACTERIOLOGY Ford. (Virulent pseudodiphtheria bacil- lus, Hamilton, Jour. Inf. Dis., /, 1904, 711 ; Ruediger's bacillus, Mellon, Jour. Bact., 2, 1917, 285; Bacillus ruedigeri Mellon, ibid., 290; Ford, Textb. Bact., 1927, 274.) From throats of fatal cases of scarlatina. Corynehactcri um segmentosum Eber- son. {Bacillus coryzae. segmentosus Cautley, Rept. Med. Officer of Health, Local Govt. Board, London, 1894-95, 455; Bacillus septus Benham, Brit. Med. Jour., ;, 1906, 1023; Eberson, Jour. Inf. Dis., 23, 1918, 17; Bacillus segmentosus Holland, Jour. Bact., 5, 1920, 220.) Rods of variable dimensions, mostly re- sembling Corynehacterium pseudodiph- iheriticum Lehmann and Neumann, but occasionally resembling Corynehacterium diphtheriae Lehmann and Neumann. Thomson and Thomson (Ann. Pickett- Thomson Res. Lab., 2, 1926, 65) do not think Cautley's bacillus is recognizable. From nasal secretions. Corynehacterium squamosum Belenky and Popova. (Cent. f. Bakt.,IAbt., Orig., 118, 1930, 444.) From normal skin of calves and small-pox vaccine. Non- hemolytic. Corynehacterium stnatum (Chester) Eberson. (Bacillus striatus flavus and Bacilhis striatus alhus von Besser, Beitr. z. path. Anat. u. Path., 6, 1888, 349; Bac- terium striatus flavus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 111; Bacterium striatum Chester, Man. Determ. Bact., 1901, 171; Bacillus flavidus Morse, Jour. Inf. Dis., 11, 1912, 281; Corynehacterium flavidum Holland, Jour. Bact., 5, 1920, 218; Eberson, Jour. Inf. Dis., 23, 1918, 5.) Eberson [loc. cit., 7) states that Bacillus diphtheroides citreus Graham-Smith (Jour. Hyg., 4, 1904, 258) corresponds with the yellow variety of this species. From nasal mucus. Resembles C or ynebacterixim seg- mentosum. Corynehacterium suis Hauduroy et al. (Le bacille pyogenes suis, Colin and Rossi, Revue gen. de Med. vetdr., 40, 1931, 137; Hauduroy etal., Diet. d. Bact. Path., 1937, 167). Causative agent of caseous suppuration of swine. Gram- negative. Corynehacterium suppuratxim Eberson. (Jour. Inf. Dis., 23, 1918, 17.) From anal pus pocket. Resembles Corynehac- terium epidermidis Eberson. Corynehacterium thermophilus Zavagli. (Amer. Jour. Hyg., 15, 1932, 504.) From raw and pasteurized milk. Grows better at 55°C than at 37°C. Corynehacterium typhi Topley and Wilson. {Bacillus typhiexanthematici Plotz, Jour. Amer. Med. Assoc, 62, 1913, 1556 ; La Presse Med., 43, 1914, 411 ; Plotz, Olitsky and Baehr, Jour. Inf. Dis., 17, 1915, 17; not Bacillus typhi exanthematici Klebs, Proc. Internat. Med. Cong., 1, 1881, 323; Corynehacterium, typhi-exan- thematici Eberson, Jour. Inf. Dis., 23, 1918, 19; Bacteri)tm typhi-exanthemalici Holland, Jour. Bact., 5, 1920, 222; Fusi- formis typhi-exanthematici Holland, ibid., 221 ; Topley and Wilson, Prin. of Bact. and Immun., 2nd ed., 1936, 349; Euhacte- rium typhi -exantheinalici Prevot, Ann. Inst. Past., 60, 1938, 295.) From blood of typhus fever patients. Corynehacterium ulcerogenes Bergey et al. {Corynehacterium diphtheriae ul- cerogenes cutaneum Mrongovius, Cent. f. Bakt., I Abt., Orig., 112, 1929, 51; Bergey et al., Manual, 4th ed., 1934, 550.) From ulcerations of the skin (human). Resemljles Corynehacterium epidermidis Eljerson and C. suppuratum Eberson. Corynehacterium vaccinae Galli-Valerio. (Cent. f. Bakt., I Abt., Orig., 36, 1904, 465.) From vaccine pustules in calves. Corynehacterium xerosis canis (Graham-Smith) Ford. {Bacillus xero- sis canis Graham-Smith, Jour. Hyg., 4, 1904, 258; Ford, Textb. Bact., 1927, 271.) From conjunctival sacs of dogs. Corynethrix hominis, C. equi, C. canis, C. anatis, etc. Czaplewski. (Deutsche med. Wchnschr., 26, 1900, 723.) Hypo- thetical species from the skin of the animals indicated. Corynethrix pseudotuberculosis murium Bongert. (Ztschr. f. Hyg., 37, 1901, FAMILY CORYXEBACTERIACEAE 40/ 472.) From a multiple, necrotic, case- ous pneumonia of mice inoculated with material from equine pneumonia. Re- garded by the author as distinct from Bacillus pseudotuberculosis muriian Kiitscher. Placed in the genus Corijne- thrix Czaplewski (Deutsche med. Wchnschr., 36, 1900, 723). Lactobacillus meleagridis Johnson and Pollard. (Diplo-bacillus Pi, Johnson and Anderson, Jour. Inf. Dis., 58, 1936, 340; Johnson and Pollard, Jour. Inf. Dis., 66, 1940, 196.) From heart, liver and yolk of moribund turkey poults. Pre- sumably a Corynebacterium , not a true Lactobacillus . Einer sporogenen Pseudo-Diphtherie- bazillus, De Simoni. (Cent. f. Bakt., I Abt., Orig., 24, 1898, 294.) From nasal secretion in ozena. Produced spores only in milk and on potato. Thought by Eberson (Jour. Inf. Dis., 23, 1918, 6) to have been a contaminated culture. Organism in M. H., De Witt. (Jour. Inf. Dis., 10, 1912, 40.) A motile, gas- producing diphtheroid isolated from a generalized diphtheria-like infection. Appendix III:* The relationships of the following soil organism are not clear, but it apparently should be placed cither in Corynebacterium or in a related genus (e.g., Mycobacterium). On agar it is rod- shaped and generally Gram-negative in young cultures, but coccoid and Gram- positive in old oidtures, a condition noted by INIellon (Jour. Bact., 2, 1917, 278) in connection with Corynebacterium enzymicum . Something similar is noted by Jensen (Proc. Linn. Soc. New So. Wales, 59, 1934, 29-62) in his description of Corynebacterium helvolum. Krassil- nikov, on the other hand (Cent. f. Bakt., II Abt., 90. 1934, 432), suggests that this species really belongs to Mycobacterium, and, after seeing a culture furnished him by Conn, has become all the more con- vinced of this relationship (personal correspondence) . Krassilnikov's studies indicate that there is a group of soil bacteria that grow as rods in young cultures with a tendency to produce branching forms in liquid media and develop coccoid bodies as they grow older. The latter then even divide and multiply like cocci. He con- siders that practically all so-called mi- crococci found among soil cultures are really the older stages of Mycobacterium spp. It is very clear that Jensen and Krassilnikov, the two leading students of the saprophytic members of this group found in soil, do not agree as to what constitutes the genus Mycobacterium; their papers appeared almost simultane- ously and clearly represent independ- ent work. Krassilnikov's description of this genus conies closest to covering organisms like the following of any of the descriptions in the literature, but it is quite different from Jensen's idea of the genus. In fact, the descriptions given by the former author seem to be more like Jensen's conception of the genus Corynebacterium. Jensen, in his description, takes into account the rela- tive acid-fast staining properties of the groups; but Krassilnikov does not men- tion either this property or the Gram stain. Inasmuch as the acid-fast prop- erty is regarded in the present classifica- tion as an important characteristic of Mycobacterium, the following species is included as an appendi.x, not of that genus, but of Corynebacterium . The re- lationships of these pleomorphic soil or- ganisms must be regarded as decidedly obscure. Lochhead (Can. Jour. Res., Sec. C, 16, 1938, 156) speaks of a Bac- terium globiforme group and Conn (Jour. Bact., 48, 1945, 359) has recently reported evidence in support of Lochhead's view- point. In all probability this group is identical in whole or in part with Krassil- nikov's Mycobacterium of soil, although the correctness of his choice of this generic name may be questioned. ♦Prepared by Prof. H. J. Conn, New York State Experiment Station, Geneva, New York, July, 1945. 408 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacterium globiforme Conn. (Conn, N. Y. Agr. Exp. Sta. Tech. Bull. 138, 1928 and 172, 1930; Cent. f. Bakt., II Abt., 76, 1928, 77; Achromobacter globi- forme Bergey et al., Manual, 3rd ed., 1930, 226.) From Latin, having the form of a globe or sphere. Short rods: 0.4 to 0.6 by 0.6 to 0.8 mi- cron, becoming coccoid in older cultures. In certain liquid synthetic media, branching forms with Gram-positive spherical granules are common. These granules have a tendency to be acid-fast. Non-motile. Rods usually Gram-nega- tive; coccoid forms usually Gram-posi- tive. Gelatin colonies: Circular, punctiform. Gelatin stab: Slow cratcriform lique- faction. Agar colonies: Circular, punctiform, translucent. Agar slant: Filiform, flat, smooth, soft, translucent, glistening growth with translucent sheen. Broth: Slight growth. Nitrites produced from nitrates in synthetic agar media. Glucose, sucrose, mannitol, and less readily lactose and various organic acids are utilized as sources of carbon and energj^ when grown in synthetic media. No visible gas production, and probably no acid except carbonic acid. Nitrogen may be obtained from am- monium sulfate, asparagine, cystine, glycerol, aspartic acid, uric acid, tyro- sin, potassium nitrate, urea and peptone. Aerobic, facultative. Optimum temperature 22°C. Source: Seventy cultures isolated from soil. Habitat: Widely distributed in soil. Genus II. Listeria Pirie.* (Listerella Pirie, Publ. So. African Inst, for Med. Res., 3, 1927, 163; not Listerella Jahn, Ber. d. deutsch. Bot. Ges., 24, 1906, 538; not Listerella Cushman, Contr. Cush- man Lab. Foram., Sharon, Mass., 9, 1933, 32; Pirie, Science, 91, 1940, 383.) Named for Joseph Lister, the English surgeon and bacteriologist. Small rods. Gram-positive. Flagellation peritrichous. Aerobic. Catalase posi- tive. Grow freely on ordinary media. Acid but no gas from glucose and a few ad- ditional carbohydrates. Pathogenic parasites. Infection characterized by a mono- cytosis. Parasitic on warm-blooded animals. The type species is Listeria monocytogenes (Murray et al.) Pirie. 1. Listeria monocytogenes (Murray et al.) Pirie. (Bacterium monocyto- genes Murray, Webb and Swann, Jour. Path, and Bact., 29, 1926, 407; Listerella hepatolytica Pirie, Publ. S. African Inst. for Med. Res., 3, 1927, 164; Listerella monocytogenes Pirie, ibid.; Listerella monocytogenes hominis Nyfeldt, Folia Haematologica, 47, 1932; Corynebacterium parvtdum Schultz, - Terry, Brice and Gebhardt, Proc. Soc. Exp. Biol. Med., 31, 1934, 1021; Pirie, Science, 91, 1940, .383; Bacillus monocytogenes Tobia, Arch, ital. med. colon., 23, 1942, 219; abst. in Cent. f. Bakt., I Abt., Ref., 144, 1943, 199.) Derived from the Greek, meaning generating monocytes. Small rods: 0.4 to 0.5 by 0.5 to 2.0 microns, with rounded ends, slightly curved in some culture media. Occur singly, in V-shaped or parallel pairs and in short chains. Motile, peritrichous (Paterson, Jour. Path, and Bact., 48, 1939, 25) with four flag6lla at ordinary temperatures with tendency toward non-motility or single flagellum at 37°C (Griffin, Jour. Bact., 48, 1944, 114). Not acid-fast. Gram-positive. Gelatin: No liquefaction. Growth is confined to the needle track. In 0.25 per cent agar, 8.0 per cent gela- tin, 1.0 per cent glucose semisolid me- dium, growth along the stab in 24 hours at 37°C, followed by irregular cloudy or * Revised by Prof. E. G. D. Murray, McGill Univ., Montreal, P. Q., Canada, Septem-ber, 1938; further revision, January, 1945. FAMILY CORYNEBACTERIACEAE 409 granular extensions into the medium; growth does not spread through the en- tire medium. This is characteristic (Seastone, Jour. Exp. Med., 62, 1935, 203). Sheep liver extract agar colonies : Cir- cular, smooth, slightly flattened, trans- parent by transmitted and milk-white by reflected light. Viscid. Sheep liver extract agar slant : Con- fluent, flat, transparent, viscid growth. Peptone agar : Growth is thinner than on liver extract agar. Blood agar : Improved growth with zone of hemolysis around colonies. Peptone broth : Surface film with flocculent sediment. Litmus milk: Slightly acid, decolor- ized. No coagulation. Glycerol-potato : Xo apparent growth. Inspissated ox servun : Grows as a very thin, transparent film. Dorset's egg medium: Very thin film. Indole not formed. Hydrogen sulfide not formed. Nitrites not produced from nitrates. Acid but no gas from glucose, rhamnose and salicin promptly, more slowly from dextrin, sucrose, soluble starch and glycerol. Acid production may be vari- able and slow from maltose and lactose. No action on arabinose, galactose, xylose, mannitol, dulcitol, inulin and inositol. All cultures give off a penetrating, rather unpleasant acid smell. Aerobic, facultative. Optimum temperature 37 ''C. Ther- mal death point 58° to 59°C in 10 minutes. Animal inoculations: Injection of rab- bits with cultures results in a very marked increase in monocytes circulat- ing in the blood. This is the most strik- ing character of the organism and is exhibited by strains derived from all sources. Infection is characterized bj' necrotic foci in various organs. Serological characters : Agglutination and absorption of agglutinin reactions show a variation in degree with different strains but there is no definite indication that strains from different kinds of ani- mal hosts are different species. Pater- son (Jour. Path, and Bact., 57, 1940, 427) concludes from his studies of the flagellar and somatic antigens of 54 cultures that four types may be recog- nized in this species. These do not bear any relation to the host species or to the geographical area from which they were isolated. Possibly related to Erysipelothrix (Barber, Jour. Path, and Bact., 48, 1939, 11). Habitat and source : Lesions in organs, blood, cerebrospinal fluid of rabbits, guinea pigs, sheep, cattle, foxes, hogs, fowls, gerbilles and man, in all of which natural disease occurs. Many cases have proved fatal. The cause of infec- tious mononucleosis in man (Nyfeldt, loc. cit.). Appendix: The following binomials have also been proposed for species in this genus. Bacterium hepatis Hiilphers. (Sven. Vet.-Tidskrift, 2, 1911, 271.) From necrosis of the liver of a rabbit. Nyfeldt (Skand. Vet.-Tidskrift, 30, 1940, 284) re- gards this as a synonym of Listcrella monocytogenes. However, failure to fer- ment lactose, rhamnose, sucrose and salicin -with fermentation of xylose, and failure to infect guinea pigs and chick- ens indicate a possible difference between the two species. Listerella hibiscus liquefaciens Naka- hama. (Jour. Agr. Chem. Soc. Japan, 16, 1940, 345.) From retted kenaf (;Hibiscus) . Listerella hominis, Listerella hovina, Listerella gallinarutn, Listerella cunicula and Listerella gerbilli Wramby. (Skand. Vet.-Tidskrift, 34, 1944, 280.) These names are given to indicate cultures of Listerella monocytogenes from man, cat- tle, chickens, rabbits and gerbilles, respectively. Listerella ovis Gill. (Australian Vet. Jour., 13, 1937, 47.) Causes circling disease of sheep. Burn (Jour. Bact., 30, 1935, 573) reports, but does not name, a new species in this genus. 410 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus III. Erysipelothrix Rosenhach* (Ztschr. f . Hyg., 63, 1909,367.) From Greek erysipelas, a disease; and thrix, hair or thread. Rod-shaped organisms with a tendency to the formation of long filaments. The filaments may also thicken and show characteristic granules. Non-motile. Gram- positive. Microaerophilic. Catalase negative. Grow freely on ordinary media. Acid but no gas from glucose and a few additional carbohydrates. Parasitic on mam- mals. The type species is Erysipelothrix rhusiopathiae (Migula) Winslow etal. 1. Erysipelothrix rhusiopathiae (Migula) Winslow et al. (Bacillus des Schweinerotlaufs, Loeffler, Arb. a. d. k. Gesundheitsamte, 1, 1886, 46; Bacillus thuillieri Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Pasteurella thuillieri DeToni and Trevisan, in Sae- cardo, Sylloge Fungorum, 8, 1889, 995; Bacillus rhusiopathiae suis Kitt, Bak- terienkunde u. path. Mikroscopie, 1893, 284; Bacterium erysipelatos suum (sic) Migula, in Engler and Prantl, Die na- tiirl. Pflanzenfam., /, la, 1895, 24; Bacterium rhusiopathiae suis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 98; Bacterium rhusiopathiae Migula, Syst. d. Bakt., 2, 1900, 43; Mycobacterium rhusiopathiae Chester, Man. Determ. Bact., 1901, 352; Erysipelothrix porci Rosenbach, Ztschr. f. Hyg., 63, 1909, 367; Winslow et al., Jour. Bact., 5, 1920, 198; Bacillus erysipelatus-suis Holland, Jour. Bact., 5, 1920, 218; Erysipelothrix erysipelatos-suis Holland, ibid. Bacillus ruboris suis Neveu-Lemaire, Precis Para- sitol. Hum., 5th ed., 1921, 24; Nocardia thuillieri Vuillemin, Encyclopedic My- cologique, Paris, 2, 1931, 125; Actino- myces thuillieri Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 45.) From Greek rhusius, reddish; pathus, a disease; red disease. Description taken in part from Karl- son, Jour. Bact., 35, 1938, 205. Slender rods: 0.2 to 0.3 by 0.5 to 1.5 microns, occurring singly and in chains. Non-motile. Gram-positive. Gelatin colonies: Hazy, bluish-gray. racemose ; situated a little below the surface, growing slowly. Gelatin stab : Small, fimbriate colonies in the stab, at times definitely arbores- cent. No surface growth. No lique- faction. Agar slant: Scant growth, translucent, moist, homogeneous. Broth: Slight turbidity, with scant, grayish sediment. Litmus milk: May become slightly acid. Indole not formed. Potato : Usually no growth. Blood serum shows scant growth. No gas from carbohydrates. Acid from glucose, galactose, fructose, lactose and more slowly from mannose and cello- biose. No acid from arabinose, xylose, rhamnose, maltose, melibiose, sucrose, trehalose, raffinose, melezitose, dextrin, starch, inulin, amygdalin, salicin, glj'cerol, erythritol, adonitol, manni- tol, sorbitol, dulcitol or inositol. Esculin not hydrolyzed. Hydrogen sulfide produced. Voges-Proskauer test negative. Methyl red test negative. Methylene blue-reduction test nega- tive. Narrow green zone of hemolysis de- velops around deep colonies on blood agar. Catalase negative. Out of 43 strains studied serologically (Watts, Jour. Path, and Bact., 60, 1940, 355), 38 appeared to be of one antigenic group, and 5 of another. * Revised by Prof. Robert S. Breed, New York State Experiment Station, August, 1938; further revision, January, 1945. FAMILY CORYXEBACTERIACEAE 411 Optimum pH 7.6. Microaerophilic. Optimum temperature 37 °C. Source : From cases of swine erysipelas. Habitat : The cause of swine erysipelas. Transmissible to gray and white mice, rabbits and pigeons. Has been trans- mitted to man by accidental inoculation. 2. Erysipelothrix muriseptica (Fliigge) Rosenbach. (Bacillus der Mausesepti- kamie, Koch, Mittheil. a. d. kaiserl. Gesundheitsamte, 1, 1881, 93; Bacillus insidiosus Trevisan, Car. di ale. nuov. gen. di Batter., 1885, 10; Bacillus muri- septicus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 250; Bacillus murimis Schroeter, in Cohn, Kryptogamen Flora V. Schlesien, 3, 1886, 162; Bacterium murisepticum Migula, in Engler and Prantl, Die natiirl. Pflanzenfam., 1, la, 1895, 24; Mycobacterium murisepticum Chester, Manual Determ. Bact., 1901, 353 ; Rosenbach, Ztschr. f . Hyg., 63, 1909, 367; Pasteurella muriseptica Bergey et al., Manual, 1st ed., 1923, 265; not Pasturella muriseptica Topley and Wil- son, Princip. Bact. and Immun., 1, 1931, 482.) From Latin jnus, muris, a mouse; Greek septicus, putrefying, septic. Rods: 0.5 by 0.8 to 1.0 micron, occur- ring singly. Xon-motile. Gram-posi- tive. Gelatin colonies: Very small, whitish, dew-like, with indefinite margin. Gelatin-stab : Filiform growth in stab, arborescent. Xo liquefaction. Agar slant : Very slight, clear, dew-like streak. Litmus milk : L'nchanged. Potato: Xo growth. Indole not formed. Xitrites not produced from nitrates. Microaerophilic. Optimum temperature 37°C. Source : From cases of mouse septi- cemia. Habitat : In fatal septicemia in white mice following injection of putrid meat infusion. Xot infectious for field mice. 3. Erysipelothrix erysipeloidis (Leh- mann and Neumann) Rosenbach. (Cladothrix des Erythema migrans, Ros- enbach, Arch. klin. Chirurg.. 36. 1887, 2; Oospora rosenbachi Sauvagais and Ra- dais, 1892, according to Brumpt, Precis de Parasit., Paris, 4th ed., 1927, 1201; Oospora erysipeloidis Lehmann and Xeumann, Bakt. Diag., 1 Aufl., 2, 1896, 392; Streptothrix rosenbachii Kruse in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 61; Actinomyces erysipeloidis Lachner-Sandoval, L^eber Strahlenpilze Strassburg, 1898, 64; Discomyces rosen- bachi Gedoelst, Champ. Paras. Homme, 1902, 177; Streptothrix erysipeloides Ca- miniti, Cent. f. Bakt., I Abt., Orig., 1907, 198; Rosenbach, Ztschr. f. Hyg., 63, 1909, 367; Nocardia rosenbachi Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 815; Babesia erysipeloides Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, ac- cording to Xannizzi, in PoUacci, Tratt. Micopat. Umana, 4, 1934, 45; Actinomyces rosenbachii Holland, Jour. Bact., 5, 1920, 216; Bacterium erysi peloidis Leh- mann and Xeumann, Bakt. Diag., 7 Aufl., 2, 1927, 499.) From Greek ery- sipelas, erysipelas; idus, shape, appear- ance. Rosenbach {loc. cit.) made a compara- tive study of the three species in this genus and came to the conclusion that they were different, although closely allied to each other. However, Rick- mann (Ztschr. f. Hyg., 64, 1909, 362) concluded that they were identical. Source : Isolated by Rosenbach (Ver- handl. d. deutsch. Gesellsch. f. Chirurg., 2, 1887, 75) in cases of human erysipe- loid. 412 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY IX. ACHROMOBACTERIACEAE BREED. (Jour. Bact,., 50, 1945, 124.) Rods, small to medium in size, cells usually uniform in shape. No branching on ordinary media, if at all. Gram-negative, rarely Gram -variable. Peritrichous or non- motile. Growth on agar slants non-chromogenic to grayish-yellow, brownish-yellow or yellow to orange. The pigment does not diffuse through the agar. Characterized by lack of power or feeble powers of attacking carbohydrates. May form acid from hexoses but no gas. May or may not reduce nitrates. May or may not liquefy gelatin. Do not liquefy agar or attack cellulose, and are not phosphorescent. Litmus milk may become faintly acid but not sufficiently acid to curdle. Usually the reaction remains unchanged or becomes alkaline. Generally salt water, fresh water and soil forms and, less commonly, parasites. Some plant pathogens may belong here. Key to the genera of familji Achromobacteriaceae. I. Non-chromogenic or at most little or no chromogenesis on agar or gelatin media. A. liitmus milk turned alkaline. No acid from carbohydrates. Genus I. Alcaligenes, p. 412. B. Litmus milk slightly acid (never curdled), unchanged or alkaline. Acid usually produced from hexose sugars. Genus II. Achromohacter, p. 417. II. Produces yellow to orange chromogenesis. A. Litmus milk slightly acid (never curdled) unchanged or alkaline. Acid usually produced from hexose sugars. Genus III. Flavobactcrinm, p. 427. Genus I. Alcaligenes Castellani and Chalmers.* (Manual Trop. Med., 3rd ed., 1919, 9.3G.) From M.L., alkali and Latin genio, to produce. Peritrichous to monotrichous, or non-motile rods. Gram-negative to Gram- variable. Do not produce acid or gas from carbohydrates. May or may not liquefy gelatin and solidified blood serum. Turn litmus milk alkaline and may or may not peptonize it. Do not form acetylmethylcarbinol. Chromogenesis when it occurs is grayish-yellow, brownish-yellow or yellow. Generally occur in the intestinal tract of vertebrates or in dairy products. The type species is Alcaligenes J aecalis Castellani and Chalmers. * Revised by Prof. H. J. Conn, New York State Experiment Station, Geneva, New York, June, 1938; further revision by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, June, 1945. FAMILY ACHROMOBACTERIACEAE 413 Key to the species of genus Alcaligenes. I. Gelatin not liquefied. A. Motile. 1. Does not produce ropiness in milk. Found in the intestinal tract. 1. Alcaligenes faecalis. 2. Produces ropiness in milk. 2. Alcaligenes viscosus. B. Non-motile. 1. Found in the intestinal tract. 3. Alcaligenes metalcaligenes . II. Gelatin liquefied. A. Motile. 1. Milk peptonized; blood serum liquefied. 4. Alcaligenes bookeri. 2. Milk not peptonized; blood serum not liquefied. .5. Alcaligenes recti. B. Non-motile. 1. Milk peptonized, slimy. 6. Alcaligenes marshallii. 1. Alcaligenes faecalis Castellani and Chalmers. (Bacillus faecalis alcalig- enes Petruschky, Cent. f. Bakt., I Abt., 19, 1896, 187; Bacterium fecalis alcalig- enes Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 73; Bacterium alcalig- enes Lehmann and Neumann, Bakt. Diag., 2 Aufl., 1899, 242; Bacillus alcalig- enes Migula, Syst. d. Bakt., 2, 1900, 737; Castellani and Chalmers, Manual Trop. Med., 1919, 936; Bacillus fecalis -alcalig- enes Holland, Jour. Bact., 5, 1920, 218; Bacterium fecalis -alcaligenes Holland, ibid.; Vibrio alcaligenes Lehmann and Neumann, Bakt. Diag., 7 Aufl., ,?, 1927, 548: Bacterium faecale alcaligenes Mo- nias. Jour. Inf. Dis., 43, 1928, 330.) From Latin /aex, dregs; M. L., fecal. Rods : 0.5 by 1.0 to 2.0 microns, occur- ring singly and in pairs, and occasionally in long chains. Motile with peritrichous flagella. In some strains, the majority of the individual cells show only a single flagellum. This is apt to be in a lateral rather than in the polar position. Gram- negative . Gelatin colonies: Circular, grayish, translucent. Gelatin stab: Gray surface growth. No liquefaction. Agar colonies : Transparent with opaque center, undulate margin. Agar slant: White, glistening, opales- cent, undulate margin. Broth: Turbid, with thin pellicle, and viscid sediment. Gives oft' ammonia. Litmus milk: Alkaline. Potato: Scanty to abundant, yellowish to brownish growth. Indole not formed. Nitrite production from nitrates vari- able. No acid or gas from carbohydrate media. No characteristic odor. Aerobic, facultative. Optimum temperature 37 °C. Source : Feces, abscesses related to intestinal tract, occasionally blood stream. Habitat: Intestinal canal. Generally considered non-pathogenic. la. Alcaligenes faecalis var. radicans Evans (Public Health Rpts., 46, 1931, 1676) is a gelatin liquefying strain. 414 MANUAL OF DETERMINATIVE BACTERIOLOGY 2. Alcaligenes viscosus (Weldiu and Levine) Weldin. {Bacillus lactis viscosus Adametz, Cent. f. i3akt., 9, 1891, 698; Bacillus viscosus lactis Kruse, in Fliigge, Die Mikroorganismen, 2, 1896, 359; Bac- terium viscosus lactis Chester, Delaware Agr. Exp. Sta. 9th Ann. Rept., 1897, 89; Bacterium lactis viscosum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 198 (Eng. ed., 1901, 196) ; Bacterium sub- viscosum Migula, Sy.st. d. Bakt., 2, 1900, 326; Group I, varieties 1, 2, 3, 4 and 5 of Harrison, Rev. G^n. du Lait, 5, 1905, 100; Bacterium visco-coccoidium Buchanan and Hammer, Iowa Agr. Exp. Stat. Re- search Bull. 22, 1915, 260; Bacillus lactis- viscosus Holland, Jour. Bact., 5, 1920, 218; Bacterium lactis-viscosus Holland, idem; Bacterium viscosum Weldin and Levine, Abst. Bact., 7, 1923, 16 (not Bacterium viscosum. Migula, Syst. d. Bakt.,^, 1900, 647) ; Lactobacillus viscosus Bergey et al., Manual, 1st ed., 1923, 244; Achromobacter viscosum Bergey et al., Manual, 2nd ed., 1925, 169; Weldin, Iowa State College Jour. Sci., 1, 1927, 186.) From Latin viscosus, viscous. Description taken largely from Long and Hammer, Iowa State Coll. Jour, of Sci., 10, 1936, 262. Rods: 0.6 to 1.0 by 0.8 to 2.6 microns, almost spherical cells frequently found, occurring singly, in pairs or short chains. Motile (Adametz, loc. cit.); non-motile (Long and Hammer, loc. cit.). Gram- negative, rarely Gram-positive. Cap- sules produced in milk cultures. Gelatin colonies: Small, gray becoming yellowish. Gelatin stab : White surface growth with villous growth in stab. Xo lique- faction. Agar colonies : After 3 to 4 days, circu- lar, 4 to 6 mm in diameter, white, viscid, shining, edge entire. Agar slant: Abundant, white, spread- ing, viscid, shining. Broth: Turbid with thin pellicle and some sediment. Ropiness generally pro- duced. Litmus milk : Ropiness produced. Pel- licle formed. Alkaline. No coagulation. Potato : Moderately heavy, dirty -white, spreading, shining growth. Indole not formed. Nitrites ordinarily not produced or produced only in a trace from nitrates. No H2S produced. Slight, if any, acid production from carbohydrates. Fat is hydrolyzed. Methyl red reaction negative. Voges-Proskauer reaction negative. Temperature relations : Growth occurs at 10° and at 20°C. At 37° and at 40°C growth variable. Aerobic. Source : Originally isolated from water. Habitat : Found in water and around dairy barns, dairy utensils. Produces ropiness in milk. Long and Hammer (Iowa State Coll. Jour. Sci., 10, 1936, 264) have described a variety of this species {Alcaligenes vis- cosus var. dissimilis) which does not produce ropiness in milk. 3. Alcaligenes metalcaligenes Castel- lani and Chalmers. (Castellani and Chalmers, Man. Trop. Med., 1919, 936; Bacterium metcdcaligcnes Weldin and Levine, Abst. Bact., 7, 1923, 13; Achro- mobacter metalcaligenes Bergey et al., Manual, 2nd ed., 1925, 169.) From Greek meta, in common with; M. L., resembling cUcaligenes. Rods : 0.6 by 1 .5 microns, with rounded ends, occurring singly and in pairs. Non- motile. Gram-negative. Gelatin stab: No liquefaction. Agar colonies : Circular, raised, smooth, amorphous, entire, gray. Agar slant : Gray, scanty, filiform, con- toured, viscid. Broth : Membranous pellicle with heavy sediment. Litmus milk: Alkaline. Potato: Scanty, glistening, smooth, sometimes faint pink. Indole not formed. FAMILY ACHROMOBACTERIACEAE 415 Nitrite production from nitrates vari- able. Starch not hydrolyzed. Blood serum not liquefied. No action on carbohydrates. Aerobic, facultative. Optimum temperature 22"C. Habitat: Intestinal canal. 4. Alcaligenes bookeri (Ford) Bergey et al. (Bacillus A of Booker, Trans. Ninth Internat. Med. Congress, 3, 1887, 598; Bacillus bookeri Ford, Studies from the Royal Victoria Hospital, Montreal, 1, 1903, 31; Bergey et al.. Manual, 1st ed., 1923, 236; Bacterium bookeri Levine and Soppeland, Eng. Exp. Sta., Iowa State College, Bui. 77, 1926, 55.) Named for the bacteriologist who first isolated this species. Rods: 0.5 by 1.5 to 2.0 microns, occur- ring singly. Motile with peritrichous flagella. Gram-negative. Gelatin colonies: Circular, brown, variable in size. Gelatin stab: Slow, saccate liquefac- tion, becoming stratiform. Agar colonies: Thin, transparent, with opaque center and indistinct margin. Agar slant : Abundant, yellowish to yel- lowish-brown . Broth: Turbid, with viscid sediment. No pellicle. Litmus milk: Alkaline. Soft curd. Litmus reduced. Peptonization. Potato: Luxuriant, yellowish-white, moist. Medium is darkened. Indole not formed. Nitrites not produced from nitrates. No acid or gas from carbohydrate media. Blood serum : Yellowish-brown growth. Gradual liquefaction. No characteristic odor. Aerobic, facultative. Optimum temperature 37 °C. Source : From alvine discharges of chil- dren suffering with cholera infantum. Habitat: Intestinal canal. 5. Alcaligenes recti (Ford) Bergey et al. {Bacterium recti Ford, Studies from the Royal Victoria Hospital, Montreal, 1, 1903, 31 ; Bergey etal., Manual, 1st ed., 1923, 236.) From Latin rectus, rectum. Rods: 0.5 by 1.5 to 2.0 microns, occur- ring singly, in pairs and in chains. Mo- tile with peritrichous flagella. Gram- negative. Gelatin colonies : Variable in size and shape, circular to oval, brown. Gelatin stab: Rapid, saccate liciuefac- tion. Agar colonies: Large, grayish-white, with opaque center. Slightly spreading. Agar slant: Grayish-white, echinulate. Broth: Turbid. No pellicle. Litmus milk : Alkaline. No peptoniza- tion. Potato: Luxuriant, moist, brownish- red. Indole not formed. Nitrites produced from nitrates. No acid or gas from carbohydrate media. Blood serum: Abundant white growth. No liquefaction. No characteristic odor. Aerobic, facultative. Optimum temperature 37°C. Source : Found but once from coecum and rectum (Ford). Habitat : Intestinal canal. 6. Alcaligenes marshallii Bergey et al. (Bacillus B of Marshall, Cent. f. Bakt., II Abt., 11, 1903, 739; Bacterium lactis marshalli Conn, Esten and Stocking, Ann. Rept. Storrs Agr. Exp. Station, 1906, 141 ; Bergey et al., Manual, 1st ed., 1923, 237.) Named for Prof. C. E. Marshall, the American bacteriologist who first isolated this species. Rods: 0.3 by 1.2 microns, occurring singly. Non-motile. Gram-negative. Gelatin colonies: Gray, granular, ir- regular, glistening. Gelatin stab: Slow, infundibuliform liquefaction. Agar slant: Filiform, gray to creamy- white, raised, becoming lemon-yellow. 416 MANUAL OP DETERMINATIVE BACTERIOLOGY Broth : Turbid, with gray ring and vis- cid sediment. Litmus milk: Alkaline, slimy, pep- tonized, strong odor. Potato : Luxuriant, lemon-yellow, smooth. Indole not formed. Nitrites not produced from nitrates. No acid or gas from carbohydrates. Aerobic, facultative. Optimum temperature 30°C. Habitat : Milk. Appendix: The following species have also sometimes been regarded as belong- ing in the genus Alcaligenes, or possess characters that indicate that they belong in this genus. Achromobacter alcaliaromaticum (Ber- lin) Bergey et al. {Bacterium alcali- aromaticum Berlin, Rev. de Microbiol, et Epidemiol., 6, 1927; Bergey et al., Man- ual, 3rd ed., 1930, 212.) From feces. See Manual, 5th ed., 1939, 509 for a de- scription of this species. This species is much like Alcaligenes faecalis. Achromobacter cxjstinovorum Barber and Burrows. (Biochem. Jour., 30, 1936, 599.) From soil. See Manual, 5th ed., 1939, 516 for a description of this species. This species is much like Alcaligenes marshallii. Achromobacter lipidis (Anderson) Allison, Anderson and Cole. {Bacter- ium lipidis Anderson, Internat. Assoc. Milk Dealers, Proc. 3()th Ann. Conven- tion, Labora ory Section, October, 1937, 19; Allison, Anderson and Cole, Jour. Bact., 36, 1938, 571.) From rancid cream. See Manual, 5th ed., 1934, 521 for a description of this species. This species is much like Alcaligenes metal- caligenes. Alcaligenes albus Bergey et al. {Bac- terium lactis album Conn, Esteu, and Stocking, Ann. Rept., Storrs Agr. Exp. Station, 1906, 143; Bergey et al.. Manual, 1st ed., 1923, 237.) From udder of cow. Gram-positive. See Manual, 5th ed., 1939, 100 for a description of this species. Alcaligenes alcalinofoetidus Hauduroy et al. {Bacillus alcalinofoetidus Cas- tellani. Jour. Trop. Med., 1930, 134; Hauduroy, Ehringer, Urbain, Guillot and Magrou, Dictionnaire des bact^ries path- ogenes. Paris, 1937, 29.) From tonsils of persons having an offensive breath. Alcaligenes ammoniagenes (Cooke and Keith) Bergey et al. {Bacterium am- moniagenes Cooke and Keith, Jour. Bact., 13, 1927, 315; Bergey et al.. Manual, 3rd ed., 1930, 367.) From feces of infants. Gram-positive. See Manual, 5th ed., 1939, 99 for a description of this species. Alcaligenes denieri Corbet. (Organ- ism No. 6, Denier and Vernet, Le Caout- chouc, 17, 1920, 10193; Quart. Jour. Rub- ber Research Inst., Malaya, 2, 1930, 152.) From the latex of Hevea brasiliensis (para rubber tree). Gram-positive. See Manual, 5th ed., 1939, 99, for a description of this species. Alcaligenes faecalis var. mariense Hau- duroy et al. {Bacillus mariense Klimenko quoted from Besson, Technique Micro- biologique, p. 904; Hauduroy et al., Diet. Path. Bact., Paris, 1937, 31.) A hydro- gen sulfide producing variety. Alcaligenes lenis De Assis. (Boletim do Inst. Vital Brasil, Niteroi, No. 14, 1930, 1.) From human blood stream. Alcaligenes stevensae Brown. (Amer. Museum Novit., No. 251, 1927, 6.) From crushed egg masses of the moth (Malacosoma americana). Said to be related to Alcaligenes bronchisepticus . Bacillus coeci Ford. (Ford, Studies from Royal Victoria Hosp., Montreal, 1, No. 5, 1903, 45.) Found in stomach and rectum of a single human subject. Much like Alcaligenes bookeri. Bacillus pylori Ford. (Ford, Studies from Royal Victoria Hosp., Montreal 1, No. 5, 1903, 44.) Found in the human stomach. Liquefied gelatin and pep- tonized casein but did not liquefy blood serum. Flavobacterium fecale Bergey et al. {Bacillus fecale aromaticum Stutzer, Cent. f. Bakt., I Abt., Orig., 91, 1923, 87; Bergey et al., Manual, 3rd ed., 1930, 150.) From feces. Resembles Alcaligenes mar- shallii. See Manual, 5th ed., 1939, 545 for a description of this species. FAMILY ACHRO.MOBACTERL^CEAE 417 Genus II. Achromobacter Bergey et al.* (Bergey et al., Manual, 1st ed., 1923, 132; Achromobacterium Richards, Proc. Soc. Agr. Bact. (British), 15th Ann. Conf., 1944, 14.) From Greek achroma, with- out color and bactrum, a staff or rod. Xon-pigment -forming (at most no pigment formed on agar or gelatin) rods. Motile with peritrichous flagella or non-motile. Gram-negative to Gram-variable. Litmus milk faintlj^ acid to unchanged or alkaline. Occur in salt to fresh water and in soil. The type species is Achromobacter liquefaciens (Eisenberg) Bergey et al. Key to the species of geiius Achromobacter. I. Motile. Flagella peritrichous. A. Gelatin liquefied. 1. Litmus milk unchanged. a. Nitrites not produced from nitrates. 1. Achromobacter liquefaciens. aa. Nitrites are produced from nitrates. 2. Achromobacter thalassius. 3. Achromobacter iophagum. 2. Litmus milk acid. a. Nitrites are produced from nitrates. 4. Achromobacter delicatulutn. B. Gelatin not liquefied. 1. Litmus milk unchanged. a. Nitrites are produced from nitrates. 5. Achromobacter aquamarinus. 6. Achromobacter cycloclastes . 2. Litmus milk slightly acid. a. Nitrites not produced from nitrates. 7. Achromobacter superficiale. II. Non-motile. A. Gelatin liquefied. 1. Litmus milk unchanged. a. Nitrites slowly produced from nitrates. 8. Achromobacter stenohalis. aa. Nitrites not produced from nitrates. 9. Achromobacter butyri. 2. Litmus milk alkaline. a. Nitrites are produced from nitrates. 10. Achromobacter stationis. B. Gelatin not liquefied. 1. Litmus milk unchanged. a. Action on nitrates not recorded. 11. Achromobacter eurydice. 2. Litmus milk acid, reduced in 5 days. a. Nitrites are produced from nitrates. 12. Achromobacter delmarvae. * Partially rearranged before his death by Prof. D. H. Bergey, Philadelphia, Penn- sylvania, September, 1937; further revision by Prof. Robert S. Breed, New York State E.xperiment Station, Geneva, New York, August, 1945. 418 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Achromobacter liquef aciens (Eisen- berg) Bergey et al. (Bacillus liquef a- ciens Eisenberg, Bakt. Diag.,3Aufl.,1891, 112; not Bacillus liquejaciens Doyen, Jour. d. connaiss. m^dic, 1889, 108; not Bacillus liquejaciens Lucet, Ann. Inst. Past., 7, 1893, 327; not Bacillus lique- faciens Migula, Syst. d. Bakt., 2, 1900, 723; Bacillus sternbergii Migula, Syst. d. Bakt., 2, 1900, 726 ; Bergey et al., Manual, 1st ed., 1923, 135.) From Latin, lique- fying. Description emended by Bergey et al . {loc. cit.). This is reported to be a com- mon water organism by Lustig (Diag. d. Bakt. des Wassers, 1893,86), by Frank- land and Frankland (Microorganisms in Water, 1894, 461) and by Horrocks (Bact. Exam, of Water, 1901, 54). Short, rather thick rods, with rounded ends, occurring singly. Motile, possess- ing peritrichous flagella. Gram-nega- tive. Gelatin colonies : Circular, gray, entire, slimy. Liquefaction. In time a putrid odor. Gelatin stab : Napiform liquefaction. Agar slant: Dirty-white, spreading growth. Broth: Turbid. Litmus milk: Unchanged. Potato: Light yellow streak. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat : Water. 2. Achromobacter thalassius ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., S, 1944, 279.) From Greek thalassius, marine, of the sea. Rods: 0.6 to 0.7 by 0.8 to 2.3 microns, with some variation in shape, occurring singly, in pairs and short chains and many cells lying side by side. Motile by means of peritrichous flagella. Gram-negative but cell walls tend to retain stain. All media except the fresh-water broth. litmus milk, and potato were prepared with sea water. Gelatin colonies: 1 mm, circular, white. Gelatin stab: Napiform liquefaction. Filiform growth along line of stab. Agar colonies: Punctiform, rough, translucent, raised. Agar slant: Moderate, glistening, beaded, watery, butyrous growth with no pigment. Sea-water broth: No pellicle, slight turbidity, scanty powdery sediment. Fresh-water broth: Fair growth. Litmus milk: No visible change. Casein not digested. Potato: No visible growth. Indole not formed. Nitrites are produced from nitrates. Does not ferment glucose, lactose, mal- tose, sucrose, xylose, mannitol, glycerol, or salicin. Starch not hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Marine bottom deposits. 3. Achromobacter iophagum (Gray and Thornton) Bergey et al. {Bacterium iophagum Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 89 ; Bergey et al., Manual, 3rd ed., 1930, 204.) From Greek ius, a poison and phagein, to eat or devour. Rods: 0.8 to 1.0 by 1.0 to 5.0 microns. Motile by means of peritrichous flagella. Gram -negative. Gelatin colonies: Quickly liquefied. Gelatin stab : Liquefied. Agar colonies: Circular or amoeboid, whitish, flat, raised, smooth, translucent, entire. Agar slant : Filiform, white to buff, flat, undulate . Broth: Turbid. Litmus milk: Unchanged. FAMILY ACHROMOBACTERIACEAE 419 Nitrites produced from nitrates. Starch hydrolyzed. Acid from glucose and sucrose. Oc- casionally from maltose and glycerol. Attacks phenol and naphthalene. Aerobic, facultative. Optimum temperature 30° to 35°C. Source : Fifteen cultures from soil. Habitat : Soil . 4. Achromobacter delicatulum (Jor- dan) Bergey et al. {Bacillus delicatulus Jordan, Report Mass. State Bd. of Health, 1890, 837; Bacterium delicatulus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 82; Bergey et al., Manual, Isted., 1923, 137.) FromLatin delicatus, soft, delicate; M. L. delicatulus, some- what delicate. Characters added to Jordan's descrip- tion by Bergey {loc. cit.) from his private notes are indicated. Steinhaus (Jour. Bact., J^, 1941, 771) apparently found the same organism and has added other characters. Rods: 1.0 by 2.0 microns, occurring singly (Jordan). Motile, possessing peri- trichous flagella. Gram-negative (Ber- gey)- Gelatin colonies: Whitish, homogene- ous, with radiate margin. Gelatin stab : Infundibuliform lique- faction. Agar slant: Whitish, glistening. Broth: Turbid, with gray pellicle and sediment. Litmus milk: Acid. Slow reduction and peptonization (Steinhaus). Potato: Thin, gray streak. Acid from glucose, sucrose, maltose and lactose (slow) (Steinhaus). No hydrolysis of starch (Steinhaus). No HjS produced (Steinhaus). Indole not formed (Bergey). Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 30° to 35°C. Source : From the effluent of a septic tank (Jordan). From water (Bergey). From the alimentary tract of an adult Colorado potato beetle (Leptinotarsa decemlineata Say) (Steinhaus) . Habitat : Presumably widely distrib- uted in nature. 5. Achromobacter aquamarinus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 264.) From Latin aqua, water, and marinus, sea. Rods: 0.8 by 1.2 to 2.0 microns, with rounded ends, occurring singly. Motile by means of a few peritrichous flagella. Gram-negative. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: 2 mm, convex, circu- lar, entire, whitish. Gelatin stab: Poor growth, no lique- faction, no pigment. Agar colonies: 2 mm, convex, smooth, circular. Agar slant: Moderate, beaded, glisten- ing, butyrous growth with no pigment. Sea-water broth: Surface ring, mod- erate turbidity, heavy viscous sediment. Fresh-water broth: Poor growth. Litmus milk: No visible change. Casein not digested. Potato: No visible growth. Indole not formed. Nitrites rapidly produced from ni- trates. Produces acid but no gas from glucose and maltose. Does not ferment lactose, sucrose, mannitol, glycerol, xylose, or salicin. Starch not hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Fats are hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Found in sea water and on sub- merged slides. Habitat : Sea water. 420 MANUAL OF DETERMINATIVE BACTERIOLOGY 6. Achromobacter cycloclastes (Gray and Thornton) Bergey et al. {Bac- terium cycloclastes Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 89; Bergey et al., Manual, 3rd ed., 1930, 212.) From Greek cyclus, ring and clastus, breaking in pieces. Rods: 1.0 to 1.5 by 1.5 to 8.0 microns. Motile with 1 to 12 peritrichous flagella. Gram-negative . Gelatin colonies: Circular, white, raised, smooth, glistening, entire. Gelatin stab: No liquefaction. Nail head growth. Agar colonies: Circular to amoeboid, white, flat to convex, smooth, glistening, translucent with opaque center, entire. Agar slant: Filiform, pale buff, raised, smooth, glistening, undulate. Broth: Turbid. Nitrites produced from nitrates. Starch not hydrolyzed. Litmus milk unchanged. No acid from carbohydrate media. Attacks phenol and naphthalene. Aerobic, facultative. Optimum temperature 30° to 35°C. Source : Three cultures from soil. Habitat : Soil . 7. Achromobacter superficiale (Jor- dan) Bergey et al. (Bacillus super- ficialis Jordan, Report Mass. State Bd. of Health, 1890, 833; Bacterium super- ficialis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 94; Bergey et al., Manual , 1 st ed . , 1 923 , 1 44 . ) From Lati ii SJiperficialis, lying on the surface. Characters added to Jordan's descrip- tion by Bergey {loc. cit.) from his private notes are indicated. Rods: 1.0 by 2.2 microns, occurring singly (Jordan). Motile, possessing peri- trichous flagella. Gram-negative (Ber- gey)- Gelatin colonies: Small, circular, gray, translucent. Gelatin stab: Scanty surface growth. Slow liquefaction. Agar slant: Limited, gray, filiform. Broth: Slightly turbid. Litmus milk: No change. Later be- coming slightly acid. Potato : No growth (Jordan). Limited growth (Bergey). Abundant (Stein- haus). Indole not formetl (Bergey). Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25° to 30°C. Source : Sewage. Gibbons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this species as occurring in the slime and feces of the cod {Gadus callarias) &nd dogfish {Squalus acanthias) . An organism apparently identical with, this organism has been found by Stein- haus (Jour. Bact., 42, 1944, 771) in the intestines of beetle larvae ( Urographus fasciata DeG.). Habitat : Presumably widely distrib- uted in nature. 8. Achromobacter stenohalis ZoBell and IJpham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 257.) From Greek stenus, narrow or close, and halinus, salty; adapted to a slight change of salinity only. Rods: 0.8 to 0.9 by 0.8 to 1.6 microns, occurring singly, in pairs and short chains. Non-motile. Capsulated. Gram -negative. All media except the fresh -water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: 1 mm, whitish, circu- lar, convex, entire. No pigment. Gelatin stab : Very slow crateriform liquefaction. Napiform in 50 days. Agar colonies: Small, circular, opales- cent, lobate edge, convex with slightly raised margin, smooth. Agar slant: Moderate, beaded, glis- tening, opalescent, beaded growth with no pigment. Sea-water broth: Moderate turbidity, viscid sediment, no pellicle or ring. Fresh-water broth: No visible growth. Litmus milk: No visible change. Casein not digested. Potato: No visible growth. FAMILY ACHROMOBACTERIACEAE 421 Indole not produced. Nitrites slowly produced from nitrates. No acid or gas from glucose, lactose, maltose, sucrose, mannitol, glycerol, xylose, or salicin. Starch not hydrolyzed. Hydrogen sulfide not produced. Ammonia produced from peptone but not from urea. Fats are not hydrolyzed. Aerobic, facultative (poor anaerobic growth). Optimum temperature 20° to 25°C. Source : Sea water, marine mud, and ■marine phytoplankton. Habitat : Sea water. 9. Achromobacter butyri Bergey et al. (Micrococcus butyri-aromafaciens Keith, The Technology Quarterly, 10, 1897, 247; Bacillus butyri aromafaciens Grimm, Cent. f. Bakt., II Abt., 8, 1902, 589; Bergey et al.. Manual, 1st ed., 1923, 148; Bacterium butyriaromafaciens Omeliansky, Jour. Bact., 8, 1923, 400.) From Latin butyrum, butter. Rods: 0.5 to 1.0 micron, nearly spheri- cal, occurring singly and in pairs. Non- motile. Gram-negative. Gelatin colonies : White, circular, smooth, glistening. Gelatin stab: White surface growth, liquefaction with white sediment. Agar slant: Abundant, white, glisten- ing. Broth : Turbid, with ring and sedi- ment. Litmus milk: Reaction luichanged. Aromatic odor. Potato: Slow and limited, white growth. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25°C. Habitat: Milk. 10. Achromobacter stationis ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 273.) From Latin statio, anchorage. Ovoid rods : 0.4 by 0.5 to 0.6 microns, occurring singly or in chains of two to three. Non-motile. Gram-positive but easily destained. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies : 0.5 to 1 mm, circular, convex, grayish-white. Gelatin stab : Very slow napiform liquefaction. Agar colonies : 1 to 2 mm, convex, lobate edge, smooth, colorless. Agar slant: Moderate, glistening, fili- form, butyrous growth with no pigment. Sea-water broth: Heavy pellicle, no turbidity, granular growtli along walls, scanty sediment. Fresh-water broth : Good growth. Litmus milk : Becomes alkaline. Casein not digested. Potato : No visible growth. Indole not formed. Nitrites rapidly produced from ni- trates. Produces acid but no gas from glucose. Does not ferment lactose, maltose, sucrose, mannitol, glycerol, xylose, or .salicin. Starch not hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Found in film of marine fouling organisms. Habitat : Sea water. 11. Achromobacter eurydice (White) Bergej' et al. [Bacterium eurydice White, U. S. Dept. of Agr., Bur. of Entomol., Circ. 157, 1912, 3 and U. S. Dept. of Agr. Bull. 810, 1920, 15; Bergey et al.. Manual, 2nd ed., 1925, 170.) From Greek Eurydice, the wife of Orpheus. Rods: Small, slender, with slightly rounded ends, occurring singly and in pairs. Non-motile. Gram-negative. 422 MANUAL OF DETERMINATIVE BACTERIOLOGY Gelatin stab : A bluish -gray growth occurs along the line of inoculation. No liquefaction. Glucose agar colonies: Bluish-gray, circular, smooth, glistening, entire. Broth: Uniform turbidity with viscid sediment. Litmus milk: Unchanged. Acid from glucose but little or no action on other carbohydrates. Potato: Slight, grayish growth. Aerobic, facultative. Innocuous when fed to bees. Not pathogenic when inoculated subcutane- ously in rabbits. Source : Occurs as a secondary invader in European foulbrood of bees. Habitat: Unknown. 12. Achromobacter delmarvae Smart. (Smart, Jour. Bact., 23, 1932, 41 and Jour. Agr. Research, 51, 1935, 363.) From Delmarva, coined from Del., Mar. and Va., the regions in which the species was found. Short rods: Average size 0.75 by 1.5 microns, with rounded ends, occurring singly, in pairs and in short chains. Non- motile. Gram-negative. Gelatin colonies : Similar to agar colonies. Gelatin stab: Scanty growth. No liquefaction. Beef-infusion agar colonies : Small, cir- cular, raised, edges smooth, glistening, translucent, bluish-white, amorphous, margin entire. Agar stab : Abundant growth. Sur- face growth round, smooth, glistening, bluish-white, raised. Filiform growth the whole length of stab, but growth best at top. Agar slant: Abundant filiform growth, raised, glistening, smooth, tran.slucent, bluish-white, no odor; old cultures slightly viscid. Medium unchanged. Nutrient broth: Turbid. Delicate white pellicle. Sediment abundant, white, slightly stringy. No odor. Color of medium unchanged. Sterile milk: Slow growth. No peptonization. Coagulation in 12 to 14 days. Milk turns chocolate brown beginning at top. Litmus milk: Acid with reduction of litmus in 5 days. Coagulation with return of pink color in 12 to 14 days. Browning of medium. Potato: Abundant growth, grayish- white, glistening, smooth, raised. Me- dium changed from white to smoke -gray. Indole not formed. Nitrites produced from nitrates in 7 days at 26°C. No HaS produced. Ammonia not formed. Diastatic action weak. Acid but no gas from glucose, lactose, glycerol and mannitol. Alkaline reac- tion and no gas from sucrose. Optimum pH 7.0. Temperature relations : Optimum 26 °C. Good growth up to 31 °C. Very slight growth at 37° and at -8°C. Facultative anaerobe. Source : Isolated from fresh straw- berries from Delaware, Maryland and Virginia. Habitat : Unknown. Appendix: Many of the following species were described before Gram and fiagella stains had been perfected. Hence it is impossible to identify them definitely as belonging to Achromobacter. Comparative study is needed in other cases before the remaining species can be placed in their proper place in the genus. Achromobacter acidum (Chester) Bergey et al. (Species No. 56 of Conn, Storrs Agr. Exper. Sta., 7th Ann. Rept. for 1894, 1895, 83; Bacterium acidum Chester, Man. Determ. Bact., 1901, 146; Bergey et al., Manual, 1st ed., 1923, 151.) From milk. See Manual, 4th ed., 1934, 246 for a description of this organism. Achromobacter agile (Ampola and Garino) Bergey et al. (Bacilhis denitri- ficans agilis Ampola and Garino, Cent. FAMILY ACHROMOBACTERIACEAE 423 f. Bakt., II Abt., 2, 1S96, 673; Bacterium denitrificans agilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 76; Bac- terium agile H. Jensen, Cent. f. Bakt., II Abt., 4, 1898, 408; Bacillus denitrifi- cans Migula, Syst. d. Bakt., 2, 1900, 796; not Bacillus denitrificans Chester, Man. Determ. Bact., 1901, 274; Bacillus agilis Chester, Man. Determ. Bact., 1901, 226; not Bacillus agilis Tschistowitsch, Berl. klin. Wchnsehr., 1892, 512; Bergey et al.. Manual, 1st ed., 1923, 138.) From cow manure. See Manual, 4th ed., 1934, 219 for a description of this organ- ism. Achromobacter album (Eisenberg) Ber- gey et al. {Bacillus albus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 171 ; Bacterium albus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 76; Bergey et al.. Manual, 1st ed., 1923, 141.) From water. Gibbons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this species from the slime on cod (Gadus callarias). See ^lanual, 4th ed., 1934, 222 for a description of this organism. Achromobacter amylovorum (Rubenl- schick) Bergey et al. {Urobacterium amylovorum Rubentschick, Cent. f. Bakt., II Abt., 64, 1925, 168; ibid., 66, 1926, 161; Bergey et al.. Manual, 3rd ed., 1930, 225.) From sewage slime. See Manual, 5th ed., 1939, 514 for a de- scription of this organism. Achromobacter anaerobium Shimwell. (Jour. Inst. Brewing, 43, 1937, 507.) From spoiled beer. Achromobacter aromafaciens (Chester) Bergey et al. (Species Xo. 41 of Conn, Storrs Agr. Exper. Sta., 7th Ann. Rept. for 1894, 1895, 57; Bacterium connii Migula, Syst. d. Bakt., 2, 1900, 440; not Bacterium connii Chester, Man. Determ. Bact., 1901, 146; Bacterium aromafaciens Chester, loc. cit., 148; Bergey et al., Manual, 1st ed., 1923, 151.) From milk sent from Uruguay to Chicago World's Fair. See Manual, 5th ed., 1939, 519 for a description of this organism. Achromobacter urcticum Rusakowa and Butkewitsch. (Microbiology (Russian), 10, 1941, 137; abst. in Cent. f. Bakt., II Abt., 105, 1942, 140.) From sea water (Barents Sea). Achromobacter candicans (Frankland and Frankland) Bergey et al. {Bacillus candicans G. and P. Frankland, Ztschr. f. Hyg., 6, 1889, 397; Bacterium candicans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 130; Bergey et al.. Manual, 1st ed., 1923, 149.) From soil. See Man- ual, 5th ed., 1939, 520 for a description of this organism. Achromobacter centropunctatum (Jen- sen) Bergey et al. {Bacterium centro- punctatus H. Jensen, Cent. f. Bakt., II Abt., 4: 1898, 410; Bacillus centropunc- fattis Chester, ]\Ian. Determ. Bact., 1901, 225; Bergey et al.. Manual, 1st ed., 1923, 139.) From cow manure. See Manual, 4th ed., 1934, 220 for a description of this organism. Achromobacter coccoideum (Chester) Bergey et al. (Species No. 16 of Conn, Storrs Agr. Exper. Sta., 6th Ann. Rept. for 1893, 1894, 51 ; Bacterium coccoideum Chester, Man. Determ. Bact., 1901, 147; Bergey et al., ■Manual, 1st ed., 1923, 152.) From ripening cream. See Manual, 5th ed., 1939, 520 for a descrip- tion of this organism. Achromobacter connii (Chester) Bergey et al. (Culture No. 55, Conn, Storrs Agr. Exp. Sta., 7th Annual Rept. for 1894, 1895, 83; Bacterium connii Chester, Man. Determ. Bact., 1901, 146; Bergey et al.. Manual, 1st ed., 1923, 149.) From milk. See Manual, 4th ed., 1934, 243 for a description of this organism. Achromobacter dendriticum (Lustig) Bergej" et al. {Bacillus dendriticus Lustig, Diagnostica dei batteri delle acque, Torino, 1890 and Diagnostik der Bakterien des Wassers, 1893, 99; Bacte- rium dendriticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103; Bergey et al.. Manual, 2nd ed., 1925, 156.) From water. See Manual, 5th ed., 1939, 504 for a description of this organ- ism. 424 MANUAL OF DETERMINATIVE BACTERIOLOGY Achromohacter epsteinii Peshkov. (Peshkov, Jour, of Biology (Russian), 6, 1937, 1003.) From water of a carp pond near Moscow. Achromohacter fermentationis (Ches- ter) Bergeyetal. {Bacterium fermenta- tionis Chester, Del. Agr. Exp. Sta. Rept. 1899, 53; Bergey et al.. Manual, 1st ed., 1923, 152.) From soil. See Manual, 4th ed., 1934, 247 for a descrip- tion of this organism. In Chester, Man. Determ. Bact., 1901 , 231 this is listed as a synonym of Bacillus foetidus-liquefaciens Tavel, Ueber Aetiol. der Strumitis, Basel, 1892. Achromohacter filefaciens (Jensen) Bergey et al. {Bacterium filefaciens H. Jensen, Cent. f. Bakt., II Abt., 4, 1898, 401; Bergey et al.. Manual, 1st ed., 1923, 153.) From dust. See Man- ual, 4th ed., 1934, 247 for a description of this organism. Achromohacter formosum (Ravenel) Bergey et al. {Bacillus formosus Rav- enel, Memoirs Nat. Acad. Sci., 8, 1896, 12; not Bacillus formosus Bredemannand Heigener, Cent. f. Bakt., II Abt., 93, 1935, 101; Bacterium formosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 91; Bergey et al., Manual, 1st ed., 1923, 136.) From soil. Gibbons (Con- trib. to Canadian Biol, and Fish., 8, No. 24, 1934, 308) reports this species from fillets of haddock {Melanogrammus aegle- finus). See Manual, 5th ed., 1939, 505 for a description of this organism. Achromohacter galophilum Bergey et al. (Culture No. 27, Baranik-Pikowsky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al.. Manual, 3rd ed., 1930, 223.) From sea water. See Manual, 5th ed., 1939, 514 for a description of this organism. Achromohacter gasoformans (Eisen- berg) Bergey et al. (Gasbildncr Bacil- lus, Tils, Zeitschr. f. Hyg., 9, 1890, 315; Bacillus gasoformans Eisenberg, Bakt. Diagnostik, 1891, 107; Bacterium gasoformans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 93 ; Bergey et al.. Manual, 1st ed., 1923, 137.) From water. See Manual, 5th ed., 1939, 503 for a description of this organism. Gas bubbles observed in plain gelatin stab. Achromohacter geminum (Chester) Ber- gey et al. {Bacillus geminus minor Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 28; Bacterium geminus minor Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 72; Bacillus geminus Chester, Man. Determ. Bact., 1901, 216; Bergey et al., Manual, 1st ed., 1923, 142.) From soil. See Manual, 5th ed., 1939, 508 for a description of this organism. Achromohacter guttatum (Zimmer- mann) Berge}' et al. {Bacillus gut- tatus Zimmermann, Bakt. unserer Trink- u. Nutzwiisser, Chemnitz, 1,^ 1890, 56; Bacterium guttatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 94; Bergey et al.. Manual, 1st ed., 1923, 140.) From water. See Manual, 5th ed., 1939, 508 for a description of this organism . Achromohacter halophilum Bergey et al. (Culture No. 36, Baranik-Pikowsky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al., Manual, 3rd ed., 1930, 220.) From sea water. See Manual, 5th ed., 1939, 513 for a description of this organism. Achromohacter hartlehii (Jensen) Bergey et al. {Bacterium hartlehii H. Jensen, Cent. f. Bakt., II Abt., 4, 1898, 449; Bacillus hartlehii Chester, Man. Determ. Bact., 1901, 226; Bergey et al., Manual, 1st ed., 1923, 139.) From soil. See Manual, 4th ed., 1934, 219 for a description of this organism. Achromohacter hyalinum (Jordan) Ber- gey et al. {Bacillus hyalinus Jordan, Report, Mass. State Bd. of Health, 1890, 835; Bacterium hyalinus Chester, Aim. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 95; Bergey et al.. Manual, 1st ed., 1923, 138.) From sand in a septic tank. See Manual, 4tli ed., 1934, 216 for a descrip- tion of this organism. Also reported by Hatcher (Jour. Elisha Mitchell Sci. Sue, FAMILY ACHROMOBACTERIACEAE 425 55, 1939, 332) from the feces of a cockroach (Periplaneta americana). Litmus milk acid and coagulated. Gram-negative. Achromohacter inu'iiclum (Pohl) Bergey et al. (Bacillus inuncius Pohl, Cent. f. Bakt.. 11, 1892, 143; Bacterium inuncius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 94; Bergey et al., Manual, 1st ed., 1923, 141.) From water. See Manual, 4th ed., 1934, 221 for a descrip- tion of this organism. Achromohacter lacticum Bergej' et al. (Kramer, Die Bakteriologie der Land- wirtschaft, 2, 1892, 24; Bergey et al., Manual, 1st ed., 1923, 152.) From slimy milk. See Manual, 5th ed., 1939, 519 for a description of this organism. This appears to refer to Loeffler's slimy milk bacillus, more correctly known as Bac- terium pituitosum Migula. Achromohacter larvae (Stutzer and Wsorow) Bergej- et al. (Enter oh acillus larvae Stutzer and Wsorow, Cent. f. Bakt., II Abt., 71, 1927, 119; Bergey et al., Manual, 3rd ed., 1930, 227.) From intestinal tract of normal and diseased caterpillars of winterwheat cutworm (Euxoa segetum). See Manual, 5th ed., 1939, 541 for a description of this organ- ism. Achromohacter liquidum (Frankland and Frankland) Berge.y et al. (Bacillus liquidus G. and P. Frankland, Ztschr. f. Hj^g., 6, 1889, 382; Bacterium liquidum Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 137; Pseudomonas liquida Chester, Man. Determ. Bact., 1901, 311; Bergey et al., Manual, 1st ed., 1923, 145.) From water. See Manual, 5th ed., 1939, 511 for a description of this organism. Achromohacter litorale (Russell) Bergey et al . ( Bacillus litoralis Russell , Ztschr. f. Hyg., 11, 1891, 199; Bacterium litoralis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 94; Pseudomonas litoralis Migula, Syst. d. Bakt., 2, 1900, 879; Bergey et al., Manual, 1st ed., 1923, 138.) See :Manual, 5th ed., 1939, 503 for a description of this organism. From mud bottom, (iulf of Xaples. Achromohacter litorale var. 2, Bois and Roy. (Xaturaliste Canadien, 71, 1945, 259.) From intestine of the codfish (Gadus callarias L.). Achromohacter middletownii (Chester) Bergey et al. (Species No. 53 of Conn, Storrs Agr. Exper. Sta., 7th Ann. Rept. for 1894, 1895, 82 ; Bacterium middletownii Chester, Man. Determ. Bact., 1901, 147; Bergey et al., Manual, 1st ed., 1923, 151.) From milk. See Manual, 4th ed., 1934, 245 for a description of this organism. Achromohacter mucidus Alford and McCleskey. (Proc . Louisiana Acad. Sci., 7, 1943, 25.) From crab meat liaving musty odor. Achromohacter nijihetsui Takeda. (Cent. f. Bakt., II Abt., 94, 1936, 48.) From fish hatchery water. Not patho- genic to salmon eggs. Achromohacter nitrovorum (Jensen) Bergey et al. (Bacterium nitrovorum H. Jensen, Cent. f. Bakt., II Abt., 4, 1898, 450; Bergey et al., :\Ianual, 1st ed., 1923, 154.) From horse manure. See Manual, 4th ed., 1934, 248 for a description of this organism. Achromohacter perolens Turner. (Aus- tralian Jour. Exp. Biol, and jSIed. Sci., 4, 1927, 57.) From musty eggs. Achromohacter pestifer (Frankland and Frankland) Bergey et al. (Bacillus pestifer G. and P. Frankland, Philosoph. Trans. Roy. Soc, London, B, 178, 1888, 277; Bacterium pestifer Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 96; Bergey et al.. Manual, 1st ed., 1923, 140.) From dust. See Manual, 5th ed., 1939, 507 for a description of this or- ganism. Achromohacter pikowskyi Bergey et al. (Culture No. 25, Baranik-Pikowsky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al., Manual, 3rd ed., 1930, 222.) From sea water. See Manual, 5th ed., 1939, 514 for a description of this organism . Achromohacter pinnatum (Ravenel) Bergey et al. (Bacillus pinnatus 426 MANUAL OF DETERMINATIVE BACTERIOLOGY Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 32; Bacterium pinnatus Chester, Ann. Rept. Del. Col. Agr. E.\p. Sta., 9, 1897, 72; Bergey et al.. Manual, 1st ed., 1923,142). From soil. See Manual, 4th ed., 1934, 223 for a description of this organism. Achromobacter ravenelii Bergey et al. {Bacillus geminus major Ravenel, Me- moirs Xat. Acad. Sci., 8, 1890,27; Bacil- lus raveneli Chester, Man. Determ. Bact., 1901, 217; Bergey et al., Manual, 1st ed., 1923, 143.) From soil. Gibbons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this species from the slime on cod (Gadus callarias). See Manual, 4th ed., 1934, 224 for a description of this organism. Achromobacter refraclans (Wright) Bergey et al. {Bacillus rejractans Wright, Mem. Nat. Acad. Sci., 7, 1894, 442; Bac- terium refraclans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 82; Bergey et al., Manual, 1st ed., 1923, 150.) From water. See Manual, 4th ed., 1934, 244 for a description of this organism. Achromobacter reticulare (Jordan) Bergey et al. {Bacillus reticularis Jordan, Rept. Mass. State Bd. of Health, 1890, 834; Bergey et al., Manual, 1st ed., 1923, 144.) From the effluent of a septic tank. See Manual, 5th ed., 1939, 610 for a description of this organism. Achromobacter rodonatum (Ravenel) Bergey et al. {Bacillus rodonatus Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 40; Bacterium rodonatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 83; Bergey et al.. Manual, 1st ed., 1923, 150.) From soil. See Manual, 4th ed., 1934, 244 for a description of this organism. Achromobacter rugosum (Chester) Bergey et al. (Species No. 27, Conn, Storrs Agr. Exp. Sta., 1893, 54; Bacillus rugosus Chester, Man. Determ. Bact., 1901 , 220; not Bacterium rugosum Henrici, Arb. Bakt. Inst. Tech. Hochschule Karlsruhe, 1, 1894, 43; not Bacillus rugosus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 438; Bacterium geminus major Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 73; Bergey et al., Manual, 1st ed., 1923, 143.) From soil. See Manual, 4th ed., 1934, 224 for a de- scription of this organism. Achromobacter sewerinii Bergey et al. (Kultur No. 3, Sewerin, Cent. f. Bakt., II Abt., 1, 1895, 162; Vibrio denitrificans Sewerin, Cent. f. Bakt., II Abt., 3, 1897, 517; Bergey et al.. Manual, 1st ed., 1923, 140.) From horse manure. Achromobacter solitarium (Ravenel) Bergey et al. {Bacillus solitarnis Rave- nel, Memoirs Nat. Acad. Sci., 8, 1896, 29; Bacterium solitarius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 71; Bergey et al., Manual, 1st ed., 1923, 143.) From soil. Gibbons (Contrib. to Cana- dian Biol, and Fish., 8, No. 22, 1934, 279) reports this species from the slime on cod {Gadus callarias). See Manual, 5th ed., 1939, 509 for a description of this or- ganism. Achromobacter stutzeri (Lehmann and Neumann) Bergey et al., {Bacillus deni- trificans II, Burri and Stutzer, Cent. f. Bakt., II Abt., 1, 1895, 392; Bacterium stutzeri Lehmann and Neumann, Bakt. Diag., 1 Aufl., ;2, 1896,237; Bacz7Z»s nitrog- encs Migula, Syst. d. Bakt., 2, 1900, 793; Bacillus stutzeri Chester, Man. Determ. Bact., 1901, 225; Bergey et al., Manual, 3rd ed., 1930, 207.) From horse manure. See Manual, 4th ed., 1934, 221 for a de- scription of this organism. Achromobacter tiogense (Wright) Ber- gey et al. {Bacillus tiogensis Wright, Memoirs Nat. Acad. Sci., 7, 1894, 441; Bacterium tiogensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 82; Bergey et al.. Manual, 1st ed., 1923, 150.) From water. See Manual, 4th ed., 1934, 244 for a description of this organism. Achromobacter ubiquitum (Jordan) Bergey et al . {Bacilhis ubiquitus Jordan, Rept." Mass. State Bd. of Health, 1890, 830; Bacterium ubiquitus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 144; Bergey et al., Manual, 1st ed., 1923, FAMILY ACHKO.MOBACTERIACKAE 427 153.) From sewage, water and dust. thias). See Alanual, 4th ed., 1934, 222 See Manual, oth ed., 1939, 517 for a de- for a description of this organism, scription of this organism. Achromohacter visco-sijmbioticum (Bu- Achromobacter venenosum (Vaughan) chanan and Hammer) Bergey et al. Bergey et al. (Bacillus venenosus {Bacillus visco-sijmhioticum Buchanan Vaughan, Amer. Jour. Med. Sci., 10^, and Hammer, Iowa Agr. Exp. Sta. Res. 1892, 191; Bergey et al., Manual, 1st ed., Bull. 22, 1915, 261 ; Escherichia symbiot- 1923, 141.) From water. Gibbons ica Bergey et al.. Manual, 1st ed., 1923, (Contrib. to Canadian Biol, and Fish., 202; Bergey et al., 3rd ed., 1930, 209.) 8, No. 22, 1934, 279) reports this species From ropy milk. See Manual, 4th ed., from the slime on cod (Gadus callarias) 1934, 223 for a description of this organ- and the feces of dogfish (Squalus acan- ism. Genus III. Flavobacterium Bergey et al.* (Bergey et al.. Manual, 1st ed., 1923, 97; Flavobacler Stewart, Jour. Mar. Biol. Assoc. Un. Kingdom, 13, 1932, 41.) From Latin^acw.s, yellow and bacterium, a small rod. Rods of medium size forming a yellow to orange pigment on culture media. Motile with peritrichous flagella or non-motile. Generally Gram-negative. Characterized by feeble powers of attacking carbohydrates, occasionally forming acid from he.xoses but no gas. Occur in water and soil. The type species is Flavobacterium aquatile (Frankland and Frankland) Bergey et al. Key to the species of genus Flavobacterium. I. Non-motile, and slow or no liquefaction of gelatin. A. Litmus milk unchanged. 1. Nitrites not produced from nitrates. 1. Flavobacterium aquatile. II. Motile with peritrichous flagella. A. Gelatin liquefied. 1. Litmus milk unchanged. a. Nitrites produced from nitrates. 2. Flavobacterium diffusum. 3. Flavobacterium okeanokoites . 4. Flavobacterium rigense. aa. Nitrites not produced from nitrates. b. From fresh water. bb. From sea water. 5. Flavobacterium devorans. 6. Flavobacterium viarinotypicum. 7. Flavobacterium marinovirosum. 8. Flavobacterium halohydrum. 9. Flavobacterium neptunium. * Partially rearranged before his death by Prof. D. H. Bergey, Philadelphia, Pennsylvania, Sept., 1937; further revision by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, June, 1945. 428 MANUAL OF DETERMINATIVE BACTERIOLOGY 2. Litmus milk alkaline. a. Nitrites produced from nitrates. 10. Flavohactcrium suaveolens. 11. Flavohacterium rhenanus. aa. Nitrites not produced from nitrates. 12. Flavohacterium marinum. 13. Flavohacterium harrisonii. B. Gelatin not liquefied. 1. Litmus milk unchanged. a. Nitrites not produced from nitrates. 14. Flavohacterium invisible. 2. Litmus milk acid. a. Nitrites not produced from nitrates. 15. Flavohacterium lactis. III. Non-motile. A. Gelatin liquefied. 1. Litmus milk unchanged. 16. Flavohncierium sewanense. 2. Litmus milk reduced. a. Nitrites not produced from nitrates. 17. Flavohacterium arhorescens. 3' Litmus milk alkaline. a. Nitrites produced from nitrates. IS. Flavohacterium. lutescens. 19. Flavohacterium fucatum. 4. Litmus milk peptonized. a. Nitrites not produced froni nitrates. 20. Flavohacterium esteroaromalicum . 5. Litmus milk acid. a. Nitrites produced from nitrates. 21. Flavohacterium, halustinum. 22. Flavohacterium dormitator . 6. Action on litmus milk not recorded. Rust-colored on blood agar. 23. Flavohacterium fcrrugineum. B. Gelatin not liquefied. 1. Litmus milk unchanged. a. Nitrites produced from nitrates. 24. Flavohacterium proteus. aa. Nitrites not produced from nitrates. 25. Flavohacterium hreve. 26. Flavohacterium solar e. C. Action on gelatin not recorded. 1. Litmus milk unchanged. a. Nitrites produced from nitrates. 27. Flavohacterium flavotenue. 1. Flavohacterium aquatile (Frank- Ztschr. f. Hyg., 6, 1889, 381; Bacterium land and Frankland) Bergey et al. aqualilis Chester, Ann. Rept. Del. Col. {Bacillus aqualilis G. imd F. Frankland, Agr. Exp. Sta., 9, 1897, 96; Bergey et FAMILY ACHROMOBACTERIACEAE 429 al., Manual, 1st ed., 1923, 100.) From Latin aquatilis, aquatic. Description taken from Frankland and Frankland and from studies by Dr. E. Windle Taylor, Metropolitan Water Board, London, on freshly isolated cultures. Rods : 0.5 by 2.5 microns, with rounded ends, occurring singly, in pairs and in chains. Oscillatory movement only ; long threads often remaining motionless (Franklands), Gram-negative (Taylor). Gelatin colonies : Center yellow-brown, with radiate arrangement of bundles of threads. Colorless margin. Very slow liquefaction (none in 6 weeks, Taylor). Gelatin stab: Yellow surface growth. Slow liquefaction. Agar slant : Yellow, smooth, glistening limited growth. Broth: Turbid with whitish sediment. No pellicle. Litmus milk : Unchanged (Taylor) . Potato: Limited, yellow streak to no growth. Indole not formed (Taylor) . Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25°C. Distinctive characters : Resembles Flavobacterium arborescens microscopi- cally; easily distinguished from this organism by its much slower and limited growth on ordinary gelatin and agar media, the marked difference in the appearance of colonies and the inability of Flavobacterium aquatile to produce more than a limited growth on potato. Source : Isolated from the water of deep wells in the chalk region of Kent, England where it occurred as a practically pure culture. Found abundantly and re- isolated by Taylor, 1941 from the same sources (personal communication). Habitat : Water. Note : The peritrichous, nitrate re- ducing and ammonia producing organism identified by Bergey {loc. cit.) in 1923, as Flavobacterium aquatile appears to have been something resembling Flavobac- terium diffusum. 2. Flavobacterium diffusum (Frank- land and Frankland) Bergey et al. {Bacillus diffusus G. and P. Frankland, Ztschr. f. Hyg., 6, 1889, 396; Bacterium diffusus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 97; Bergey et al., Manual, 1st ed., 1923, 100.) From Latin diffusus, spreading out, diffuse. Description completed from Harrison (Canadian Jour. Res., 1, 1929, 233) as indicated. Rods: 0.5 by 1.5 microns, occurring singly and in chains. Motile, possessing peritrichous flagella. Gram-negative (Harrison) . Gelatin colonies: Thin, bluish-green, spreading, later faint yellow. Gelatin stab : Thin, glistening, yellow'- ish-green surface growth. Slow crateri- form liquefaction. Agar slant : Thin, light yellow, glisten- ing. Broth: Turbid, with greenish-yellow sediment. Litmus milk: Unchanged (Harrison). Potato : Thin, smooth, greenish-yellow, glistening growth. Indole not formed (Harrison). Nitrites produced from nitrates (Har- rison). Slight acidity from glucose. No acid from sucrose and lactose (Harrison). Aerobic, facultative. Optimum temperature 25° to 30°C. Source: Originally found in soil. Found also by Tataroff (Die Dorpater Wasserbakterien, Dorpat, 1891, 58) in fresh water and by Harrison (loc. cit.) from skin of halibut from both the At- lantic and Pacific shores of Canada. Habitat: Soil, fresh and sea waters. 3. Flavobacterium okeanokoites Zo- Bell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 270.) From Greek Oceanus, the ocean god, the ocean and coites, bed. Rods: 0.8 to 0.9 by 1.2 to 1.6 microns, with rounded ends, many coccoid, oc- curring singly and in long chains. 430 MANUAL OF DETERMINATIVE BACTERIOLOGY Motile by means of peritrichous flagella. Gram-negative. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Small, circular, con- vex, entire, rust or orange colored, digest gelatin. Gelatin stab: Slow napiform liquefac- tion, yellow growth. Agar colonies : 2 mm, circular, entire, smooth, convex. Agar slant: Moderate, filiform, glisten- ing, butyrous growth with yellow pig- ment. Sea -water broth : No pellicle, moderate turbidity, moderate viscid sediment. Fresh-water broth: Good growth. Litmus milk: No visible change. Casein is digested. Potato: No visible growth. Indole not formed. Nitrites slowly produced from nitrates. Does not produce acid or gas from glucose, lactose, maltose, sucrose, glyc- erol, mannitol, xylose, or salicin. Starch not hydrolyzed. Hydrogen sulfide is formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source: Marine mud. Habitat: Sea water. 4. Flavobacterium rigense Bergey et al. {Bacillus brunneus rigensis Bazarew- ski, Cent. f. Bakt., II Abt., 15, 1905, 1; Bergey et al., Manual, 1st ed., 1923, 100.) From Riga, the name of the city where the species was isolated. Rods : 0.75 by 1.7 to 2.5 microns, occur- ring singly. Motile, possessing peri- trichous flagella. Gram -negative. Gelatin colonies : Circular, entire to undulate, grayish-white, homogeneous. Gelatin stab: Smooth, yellowish sur- face growth. Infundibuliform liquefac- tion. Brownish-yellow sediment. Agar slant : Narrow, whitish streak, be- coming yellowish -brown, spreading. Pigment is water and alcohol soluble. In- soluble in ether. Broth : Turbid with pellicle and brown- ish sediment. Cells capsulated. Litmus milk: Unchanged. Potato: Yellow, spreading growth. The growth turns brownish. Hydrogen sulfide not formed. Indole not formed. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 30°C. Brown- ish colors develop best at lower tempera- tures. Orange-yellow colors develop best at 37°C. Plabitat: Soil. 5. Flavobacterium devorans (Zimmer- mann) Bergey et al. {Bacillus devorans Zimmermann, Bakt. unserer Trink- u. Nutzwiisser, Chemnitz, 1, 1890, 48; Bacterium devorans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 96; Bergey et al.. Manual, 1st ed., 1923, 102.) From Latin devorans, devouring. Characters added to Zimmermann's description by Bergey {loc. cit.) from his private notes are indicated. Steinhaus (Jour. Bact., 42, 1941, 771) apparently found the same organism. Rods : 0.7 by 0.9 to 1.2 microns, occur- ring singly, in pairs and chains. Motile (Zimmermann), possessing peritrichous flagella (Bergey). Gram-negative (Zim- mermann). Gelatin colonies : Circular, white, gran- ular to filamentous, becoming yellowish- gray. Gelatin stab: Slow infundibuliform liquefaction. Agar slant: Thin, gray, spreading. Broth: Turbid. Litmus milk: Unchanged. Potato: No growth (Zimmermann). Yellowish -gray streak (Bergey). Indole not formed. Nitrites not produced from nitrates (Bergey). FAMILY ACHROMOBACTERIACEAE 431 Aerobic, facultative. Optimum temperature 25° to 30°C. Source : From water at Chemnitz (Zim- mermann). From water (Bergey). From alimentary tract of the nine-spotted lady beetle {Coccinella novemnotata Habst.) (Steinhaus). Habitat : Water. 6. Flavobacterium marinotypicum Zo- Bell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 268.) From Latin jnan?) MS, of the sea and lypi- cus, typical. Rods: 0.5 to 0.7 by 1.4 to 2.0 microns, occurring almost entirely as single cells. Motile by means of four or more peri- trichous flagella. Gram-negative. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Very minute, yellow, with slow liquefaction. Gelatin stab : Crateriform liquefaction t)ecoming stratiform. Filiform along line of stab. Agar colonies : Minute, circular, entire, convex, yellow. Agar slant: Scanty, filiform, butyrous, shiny growth with yellow pigment. Sea-water broth: Scanty, yellowish pellicle, heavy turbidity, slight viscid sediment . Fresh-water broth : Good growth. Litmus milk: Decolorized, neutral, greenish pellicle, slow peptonization. Potato: Abundant, shiny, greenish-yel- low growth. Potato darkened. Indole not formed. Nitrites not produced from nitrates. Produces acid but no gas from glucose and glycerol. Does not ferment lactose, sucrose, mannitol, xylose, or salicin. Starch not hydrolyzed. Hydrogen sulfide is formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. .\erobic, facultative. Optimum temperature 20° to 25°C. Source : Sea water and marine mud. Habitat : Sea water. 7. Flavobacterium marinovirosum Zo- Bell and L^pham. (Bull. Scripps. Inst, of Oceanography, Univ. Calif., 5, 1944, 271.) From Latin marinus, of the sea, and virosus, covered with slimy liquid or ooze. Rods: 0.7 to 0.8 by 0.8 to 2.8 microns, with rounded ends, occurring singly and in long chains. Motile by means of peri- trichous flagella. Gram-negative. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Small, circular, raised, rust-colored. Slowly digest gel- atin. Gelatin stab : Crateriform liquefaction becoming stratiform. Light orange pig- ment. Agar colonies : 1 to 2 mm, circular, con- vex, entire, smooth. Agar slant: Moderate, filiform, glisten- ing, mucoid growth with grayish-yellow pigment . Sea -water broth: Heavy turbidity, no pellicle, abundant viscid sediment. Fresh-water broth: Good growth. Litmus milk: No visible change. Casein is digested. Potato: No visible growth. Indole not formed. Nitrites not produced from nitrates. Does not ferment glycerol, glucose, lactose, maltose, sucrose, mannitol, xylose, or salicin. Starch not hydrolyzed. Hydrogen sulfide is formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Sea water and marine mud. Habitat: Sea water. 8. Flavobacterium halohydrixxm ZoBell and Upham. (Bull. Scripps Inst, of 432 MANUAL OF DETERMINATIVE BACTERIOLOGY Oceanography, Univ. Calif., 5, 1944, 278.) From Greek hals, salt and hydror, water. Short rods: 0.6 by 0.8 to 1.0 microns, occurring singly. Motile by means of many peritrichous flagella. Gram-nega- tive. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Small, circular, orange. Gelatin stab: Napiform liquefaction becoming crateriform. Beaded along line of stab. Agar colonies: 2 mm, pulvinate, circu- lar, entire, smooth. Agar slant : Moderate, glistening, echin- ulate, butyrous growth with yellow pig- ment. Sea-water broth: Yellow surface ring, heavy turbidity, moderate viscid sedi- ment. Fresh -water broth: No visible growth. Litmus milk: No visible change. Casein not digested. Very poorly tolerant of increases or de- creases in salinity. Potato: No visible growth. Indole not formed. Nitrites not produced from nitrates. Produces acid but no gas from glucose, lactose, maltose, sucrose, and salicin. Does not ferment glycerol, mannitol, or xylose. Starch is hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Sea water and marine mud. Habitat : Sea water. 9. Flavobacterium neptunixun ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 278.) From Latin Neptunius, god of the sea. Rods: 0.5 to 0.6 by 1.6 to 4.5 microns, many bent rods, occurring singly and in short chains. Motile by means of long, peritrichous flagella. Gram-negative. All media except the fresh-water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Small, circular, darker centers, sink in gelatin, faintly yellow. Gelatin stab : Slow napiform liquefac- tion. Filiform growth along line of stab. Agar colonies: 2 mm, circular, smooth, entire, convex, dark centers with buff pigment. Agar slant : Luxuriant, echinulate, glis- tening, slightly mucoid growth with buff to yellow pigment. Agar discolored brown. Sea-water broth: Heavy pellicle, scanty turbidity, scanty sediment. Fresh-water broth : No visible growth. Litmus milk: No visible change. Casein not digested. Potato : No visible growth. Indole not formed. Nitrites not produced from nitrates. Produces acid but no gas from glucose, lactose, maltose, and salicin. Does not ferment glycerol, mannitol, xylose, or sucrose. Starch is hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Marine bottom deposits. Habitat : Sea water. 10. Flavobacterium suaveolens Soppe- land. (Jour. Agr. Res., 28, 1924, 275.) From Latin siiaveolens, of a sweet odor. Rods: 0.6 to 0.8 by 1.0 to 1.2 microns, with rounded ends, occurring singly and in pairs. Motile, with peritrichous fla- gella. Gram-negative on plain agar. Gram-positive in young culture on milk powder agar. Gelatin stab: Rapid stratiform lique- faction. Medium becomes brown. FAMILY ACHROMOBACTERIACEAE 433 Agar colonies: Small, circular, smooth, yellow, amorphous, undulate margin. Agar slant: Moderate, flat, glistening, opaque, butyrous, yellow, with aromatic odor. Broth: Turbid with scanty sediment. Aromatic odor, becoming cheesy. Litmus milk: Peptonized. Alkaline. Potato: Abundant, yellow, glistening, becoming brown. Indole formed. Nitrites are produced from nitrates. Hydrogen sulfide formed. Slight acid but no gas from glucose, sucrose and glycerol. No acid from lac- tose . Starch hydrolyzed. Blood serum is liquefied. Aerobic, facultative. Optimum temperature 25°C. Source : Dairy wastes. Habitat: Unknown. 11. Flavobacterium rhenanus (Migula) Bergey et al. (Rhine water bacillus of Burri, Frankland and Frankland, Micro- organisms in Water, 1894, 483; Bacillus rhenanus Migula, Syst. d. Bakt., 2, 1900, 713; Bacillus rheni Chester, Manual De- term. Bact., 1901, 251; Bergey et al.. Manual, 1st ed., 1923, 103.) Named for the Rhine River. Characters added to Burri's descrip- tion by Bergey (loc. cit.) from his private notes are indicated. Steinhaus (Jour. Bact., ^2, 1941, 771) apparently found the same organism and has added other char- acters. Rods: 0.7 by 2.5 to 3.5 microns, with rounded ends, occurring singly and in chains (Burri). Motile, possessing peri - trichous flagella (Bergey). Gram-nega- tive (Bergey). Gelatin colonies: Convex, colorless, transparent, becoming yellowish. Gelatin stab: Infundibuliform lique- faction. Agar colonies : Small, smooth, convex, entire. Glycerol agar slant: Thin, shining, honey-colored. Growth dry and tough. Broth: Turbid, with orange-colored pellicle and sediment. Litmus milk: Soft coagulum, becoming slightly alkaline with yellow ring. Potato: Moist, glistening, thin, flat, orange to rust-colored. Indole not formed (Bergey). Nitrites produced from nitrates (Ber- gey). Acid from glucose, maltose, and su- crose but not lactose (Steinhaus). No hydrolysis of starch (Steinhaus). No HoS produced (Steinhaus). Aerobic facultative. Optimum temperature 30°C. Source : From Rhine River water (Burri). From water (Bergey). From eggs in ovary of a walking stick (DiapheromerafemorataSsiy) (Steinhaus). Habitat : Presumably widely distrib- uted in nature. 12. Flavobacterium marinum Harri- son. (Canadian Jour, of Research, /, 1929, 234.) From Latin marinits, per- taining to the sea. Rods: 0.8 by 1.2 to 1.3 microns, with rounded ends. Occur singly and in pairs. Motile with 4 to 5 peritrichous flagella. Encapsulated. Gram-variable. Show blue granules in Gram-negative rods . Gelatin colonies : Circular, iridescent, whitish margin with pale yellow center. Liquefaction. Gelatin stab : Saccate to stratiform liquefaction. Agar colonies: Circular, pale yellow, smooth, convex, granular, reticulate edge. Agar slant: Amber-yellow, slightly raised, spreading, smooth, glistening, transparent. Ammonium phosphate agar : Scant growth. Broth : Turbid, sediment. Litmus milk: Alkaline. Digestion without coagulation. Clear serum. Potato: Abundant, amber-yellow, be- coming dirty yellow, spreading, glisten- ing. 434 MANUAL OF DETERMINATIVE BACTERIOLOGY Indole not formed. Nitrites not produced from nitrates. Trace of ammonia formed. Faint acidity from glucose. No action on lactose or sucrose. Loeffler's blood serum not liquefied. Faint yellow spreading gi'owth. No H2S formed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Isolated from living halibut obtained at 30 to 50 fathoms, Pacific Ocean. Gibbons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) re- ports this species as occurring in the slime and feces of cod (Gadus callarias), hali- but (Ilippoglossus hippogloss^is) and skate {Raja erinacea) . Habitat : Skin and feces of fishes. 13. Flavobacterium harrisonii Bergey et al. (Variety No. 6, Harrison, Rev. gen. du Lait, 5, 1905, 129; Bacillus lactis harrisonii Conn, Esten and Stocking, Ann. Rept. Storrs Agr. Exp. Sta., 1906, 169; Bergey et al.. Manual, 1st ed., 1923, 104.) Named for Prof. F. C. Harrison, the Canadian bacteriologist who first iso- lated this species. Rods : 0.25 to 0.75 by 0.3 to 3.5 microns, occurring singly and occasionally in short chains. Motile, possessing peritrichous flagella. Gram-negative. Gelatin colonies: Small, gray, glisten- ing, lobular, citron-yellow, slimy. Gelatin stab: Villous growth in stab. Slow crateriform to napiform liquefac- tion. Agar slant : Lu.xuriant, viscous, spread- ing, becoming dirt}', to brownish citron- yellow. Broth : Turbid, with viscid ring and gelatinous sediment, sweetish odor, al- kaline. Litmus milk : Colorless to gray and slimy, becoming yellow, alkaline. Potato : Luxuriant, yellow, spreading, slimy. Indole not formed. Glucose, lactose, maltose and sucrose broth turn alkaline with a disagreeable odor. Reaction of glycerol broth remains neutral. Aerobic, facultative. Optimum temperature 25°C. Source : Slimy milk. Habitat : Unknown. 14. Flavobacterium invisibile (Vaughan) Bergey et al. (Bacillus invisihilis Vaughan, American Jour. Med. Sci., IO4, 1892, 191 ; Bacterium invisibilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 77; Bergey et al.. Manual, 1st ed., 1923, 109.) From Latin invisibilis, not visibile. Rods: 0.6 to 0.7 by 1.2 to 2.0 microns, occurring singly. Motile, possessing peritrichous flagella. Gram-negative. Gelatin colonies : Pale yellow, burr- like, with irregular margin. Gelatin stab : Scanty growth on surface. Good growth in stab. No liquefaction. Agar colonies: White, convex, smooth, serrate. Agar slant : Limited, thick, white streak. Broth: Turbid. Litmus milk: Unchanged. Potato : No growth. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 35°C. Habitat : Water. 15. Flavobacterium lactis Bergey et al. {Bacillus uruiiiaLicus lactis Grimm, Cent. f. Bakt., II Abt., 8, 1902, 584; Bacillus aromaticus Grimm, ibid., 589; not Bacillus aruinaticus Pammel, Bull. 21, Iowa Agr. Exp. Sta., 1893, 792; Bergey et al., Manual, 1st ed., 1923, 108.) From Latin lac, milk. Rods: 0.7 to 1.0 by 3.5 to 4.0 microns, occurring singly, in pairs and in chains. Motile, possessing peritrichous flagella. Gram -negative. Gelatin colonies ; Circular, light yellow, slimy. Concentrically ringed, undulate. Gelatin stab: Slimy surface growth. No liquefaction. FAMILY ACHROMOBACTERIACEAE 435 Agar slant: Slimj-, yellowish, smooth, moist. Broth : Turbid, with abundant sedi- ment. Litmus milk : Slightly acid. Potato : Thick, slimy, brownish, with yellowish margin. Indole not formed. Nitrites not produced from nitrates. Cultures have pleasant odor. Aerobic, facultative. Optimum temperature 25°C. Source : Isolated from milk. Habitat : Unknown. 16. Flavobacterium sewanense (Ka- lantarian and Petrossian) Bergey et al.- (Bacterium sewanense Kalantarian and Petrossian, Cent. f. Bakt., II Abt., 85, 1932, 431 ; Bergey et al., IManual, 4th ed., 1934, 160.) From M. L., Sevan, a lake in Armenia. Straight or curved rods: 1.0 to 2.0 b\' 4.0 to 5.0 microns on Molisch's agar; on meat extract agar and potato agar they are short or even coccoid. Ends rounded, occurring singly or in pairs. Non-motile. Gram reaction not given. Presumably negative . Gelatin stab : Slow liquefaction. Agar colonies: Circular, raised, glisten- ing, dirty white. Deep colonies yellow and lens-shaped. Agar slant : Abundant, dirty yellow, glistening, raised. Broth : Turbid with characteristic growth forms. Pellicle formed in old cul- tures. Milk: Unchanged. Potato : Yellow, raised, glistening, with darkening of the medium. No visible gas produced from carbo- hydrates. Crystals of calcium carbonate form in old cultures on CaCU and Molisch's agar. Aerobic, facultative. Optimum temperature 20°C. Source : Isolated from pellicle formed on surface of fish infusions in Lake Sevan and tap waters containing 1 per cent CaCL. Habitat : Sea water. Thought to pro- duce deposits of CaCOa in Lake Sevan, S. S. R. Armenia. 17. Flavobacterium arborescens (Frankland and Frankland) Bergey et al. {Bacillus arborescens Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 379; also see Tils, Ztschr. f. Hyg., 9, 1890, 312; Zimmermann, Bakt. unserer Trink- u. Nutzwasser, 2, 1894, 20; and Wright, Mem. Nat. Acad. Sci., 7, 1894, 446, var. a and b; Bacterium arborescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 106; IMigula, Syst. d. Bakt., 2, 1900, 468 uses Bacillus arborescens in the text by mistake as Bacterium is used for other species in the genus and Bacterium ar- borescens is used in the index, p. 1058; not Bacillus arborescens Chester, Man. Determ. Bact., 1901, 249; Erythrobacillus arborescens Holland, Jour. Bact., 5, 1920, 217; Bergey et al., Manual, 1st ed., 1923, 113.) From Latin arborescens, becoming a tree or tree-like. Rods: 0.5 by 2.5 microns, occurring singly and in chains. Non-motile (Frank- lands). Gram-negative (Zimmermann). Gelatin colonies : Radiate branching filaments. Center yellowish, border translucent. Gelatin stab : Liquefied with yellow deposit. Agar slant : Dirty orange growth. Broth: Turbid, with orange sediment. No pellicle. Litmus milk : Slow coagulation ; litmus reduced. Reaction unchanged (Wright). Potato : Deep orange, luxuriant growth. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30°C. May belong to Corynebacterium (Leh- mann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 709). Source : From river and lake water. Habitat: Water. 17a. Bacillus arborescens Chester. (Bacillus arborescens non-liquefaciens Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 436 MANUAL OF DETERMINATIVE BACTERIOLOGY 39; Bacterium arborescens non-Iique- faciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103; not Bac- terium arborescens non-liquefaciens von Rigler, Hyg. Rund., 12, 1902, 479; Ches- ter, Man. Determ. Bact., 1901, 249.) Regarded by author as a non-liquefying strain of Bacillus arborescens Frankland and Frankland. Not a yellow chronio- gen. From soil. 18. Flavobacterium lutescens (Mig- ula) Bergey et al. (Der gelbe Bacillus, Lustig, Diagnostik der Bakterien des Wassers, 1893, 78; Bacterium lutescens Migula, Syst. d. Bakt., 2, 1900, 476; Bergey et al., Manual, 1st ed., 1923, 114.) From Latin lutum, yellow; lutescens, be- coming yellowish. Rods: 0.5 by 0.95 micron, occurring singly and in pairs. Non-motile. Gram- negative. Gelatin colonies: Circular, yellow, lobate. Gelatin stab : Slow liquefaction. Agar slant : Pale yellow, becoming golden yellow. Broth: Turbid. Litmus milk: Alkaline. Potato: Lu.xuriant, golden-yellow growth. Indole not formed. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 30° to 35°C. Source : From water. Gibbons (Con- trib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this species as oc- curring in the slime of the cod (Gadus callarias ) . Habitat : Fresh and salt water. 19. Flavobacterium fucatiun Harrison. (Canadian Jour, of Research, 1, 1929, 232.) From Latin /(tea/ us, painted, col- ored. Rods: 0.8 to 1.0 by 2.5 to 3.5 microns, slightly bent, with rounded ends. Gran- ular with diphtheroid forms at 37°C. Non- motile. Gram-negative. Gelatin colonies : Circular, yellow, en- tire, paler at edges. Gelatin stab : Crateriform liquefaction. Agar colonies: Circular, buff -yellow, smooth, shiny, convex to pulvinate, gran- ular, entire. Agar slant : Moderate, light buff -yellow, spreading, shiny, smooth. Ammonium phosphate agar : Good growth in 6 days. Broth : Turbid, becoming clear, pellicle and yellow sediment. Litmus milk: Alkaline. Peptonized. Clear serum. Yellow sediment. Potato : Abundant, pale buff-yellow, smooth, spreading, becoming orange-yel- low. Indole not formed. Nitrites produced from nitrates. Traces of ammonia formed. No acid from glucose, lactose or suc- rose. Loeffler's blood serum not liquefied. Light buff-yellow growth becoming ochra- ceus salmon. No H2S formed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Repeatedly isolated from living halibut obtained at 30 to 50 fathoms, Pacific Ocean. Also isolated by Gibbons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) from cod (Gadus callarias) and dogfish {Squalus acan- thias). Habitat : Skin of fishes. 20. Flavobacterium esteroaromaticxmi (Omelianski) Bergey et al. (Bacterium esteroaromaticum Omelianski, Jour. Bact., 8, 1923, 407; Bergey et al., Manual, 3rd ed., 1930, 149.) From M. L., ester and Greek aromaticus, aromatic. Rods: 0.5 by 1.0 to 3.0 microns. Non- motile. Gram reaction not recorded. Gelatin stab: Crateriform liquefaction with odor of musk melons. Agar colonies: Circular, yellow-brown, with fimbriate margin and a fruity aroma. FAMILY ACHKO.MOBACTEKIACEAE 43; Broth : Turbid, slight sediment . Faint fruity aroma. Litmus milk: Peptonized. Cheesy odor. Potato: Abundant growth. Disagree- able odor. Loeffler's blood serum: Liquefied. Indole not formed. Nitrites not produced from nitrates. Ammonia formed. Hydrogen sulfide formed. Fat hj^drolyzed. Methylene blue reduced. No acid from carbohj-drates. Aerobic, facultative. Optimum temperature 30°C. Source : Accidental contaminant in rab- bit brain containing rabies virus. Habitat : Presumably widely distrib- uted . 2L Flavobacterium balustinum Harri- son. (Canadian Jour. Research, /. 1929, 234.) Rods : 0.6 by 2.0 to 4.0 microns, forming short chains. Non-motile. Gram -nega- tive. Gelatin colonies: Circular, bright yel- low center, entire. Gelatin stab: Liquefied. Agar colonies: Punctiform, cadmium- yellow, convex, shiny, transparent. Agar slant : Egg yolk-yellow, semi- transparent streak, smooth, shiny, be- coming brownish -yellow. Ammonium phosphate agar : Slight yel- low growth. Broth: Turbid, with yellow sediment. Litmus milk : Slightly acid with yellow SBdiment. Potato : Scant, yellow growth. Indole not formed. Nitrites (trace) produced from ni- trates. Ammonia not formed. Faint acidity from glucose. No action on lactose or sucrose. Loeffler's blood serum not liquefied. Egg yolk-like growth. No H2S formed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Isolated from living halibut obtained at 30 to 50 fathoms. Pacific Ocean. Habitat : Skin of fishes. 22. Flavobacterium dormitator (Wright) Bergey et al. {Bacillus dormi- tator Wright, Memoirs Nat. Acad. Sci., 7, 1895, 442; Bacterium dormitator Chester, Ann. Rept. Del. Agr. Exp.'Sta., 9, 1897, 109; Bergey et al., Manual, 1st ed., 1923, 115.) From Latin dormitator, one who sleeps. Description completed from Harrison (Canadian Jour. Res., 1, 1929, 233) whose cultures differed in some particulars from Wright's. Rods with conical ends, occurring singly, in pairs and in chains. Non-mo- tile. Gram-negative (Harrison). Gelatin colonies : Small, yellow, slightly granular, liquefying. Gelatin stab: Infundibuliform lique- faction, yellow sediment. Agar slant : Yellow, glistening, translu- cent . Ammonium phosphate agar: Slight yel- low growth. Broth: Turbid, with slight pellicle and yellow sediment. Litmus milk: Slightly acid; litmus re- duced. Harrison reports no reduction. Potato: Slight, transparent, yellow growth. Indole not formed (Harrison). Nitrite (trace) produced from nitrates (Harrison). Acid from glucose, sucrose, glycerol and mannitol. No acid from lactose, raffinose, and inulin (Harrison). Aerobic, facultative. Optimum temperature 30^C. Source : Originally isolated from fresh water at Philadelphia. Later isolated by Harrison (loc. cit.) from skin of halibut taken in Pacific ocean off Canada. Gib- bons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this 438 MANUAL OF DETERMINATIVE BACTERIOLOGY species as occurring in the slime of a had- dock {Melanogramrnus aeglefinus). Habitat : Fresh and salt water. 23. Flavobacterium ferrugineum Sick- lesand Shaw. (Jour. Bact., 28, 1934, 421.) From Latin f err ugineus, resembling rust. Small, slender rods: Less than 0.5 by 0.7 to 1.0 micron, occurring singly and in pairs. Non-motile. Gram-negative. Gelatin : Liquefaction in one week at 37°C ; at room temperature liquefaction slower, napiform ; yellow sediment along line of puncture. Blood agar colonies: Dull, rust-colored, 1 mm in diameter, round, entire, umbili- cate, rather dry. Agar colonies: Similar to blood agar colonies but yellowish-gray in color. Blood agar slants: Moderate growth, rust-colored, rather drj'. Agar slants: Growth very slight, thin, yellowish-gray. Beef -infusion broth: No growth. Beef extract broth : Moderate even tur- bidity. Adding type -specific carbohy- drate results in a heavier growth with yellow sediment. Potato : Moderate growth, bright orange in color. Potato darkened. Very active hydrolysis of starch. Acid but no gas from glucose, lactose, sucrose, maltose, dextrin and inulin ; very slight action on mannitol; no action on salicin. Limits of growth: Optimum pH 7.0 to 7.5. Minimum 6.5. Maximum 9.0. Temperature relations: Optimum 35° to 37 °C. Minimum 22°C. Maximum 39°C. Thermal death point 52°C for 10 minutes. Enzyme produced by strain against pneumococcus carbohydrate with- stands 56°C for 10 minutes. Facultative aerobe. Distinctive character : Decomposes the non-type-specific carbohydrate obtained from a degraded type I pneumococcus. Source: Several, strains isolated from swamps and other uncultivated soils. Habitat: Soil. 24. Flavobacterium proteus Shimwell and Grimes. (Bacterium Y, Shimwell and Grimes, Jour. Inst. Brewing, 4-3, N.S. 23, 19.36, 119 ; Shimwell and Grimes, ibid., 348.) From Latin Proteus, a sea god who often changed in form. Rods: 0.8 to 1.2 by 1.5 to 4.0 microns, occurring singly or in chains, and having rounded ends. Highly pleomorphic. Thickened filaments and spindle-shaped swellings common . Probably non-motile . Gram-negative. Wort-gelatin plate : Surface colonies irregular, up to 1 mm in diameter, gray- ish-white or yellowish, fiat or slightly raised, margin entire to lobate or crenate. Deep colonies circular, small, yellowish. Wort-gelatin streak: Scanty, filiform or beaded, .slightly raised, at first almost transparent, later more opaque and whit- ish-buff. Wort-gelatin stab : Scanty, filiform or beaded, almost colorless. No liquefac- tion. Wort-agar plate: Colonies small, pale, buff-colored, resembling bread-crumbs in shape . Wort-agar streak : Similar to wort-gela- tin streak. Sometimes a slight metallic sheen on old cultures. Broth : Turbid in 24 hours at 30°C, with a slight surface scum. Litmus milk : Unchanged. Potato: A slight, barely visible growth consisting of a narrow filiform dirty yel- low line. Indole not produced. Nitrites are produced from nitrates. Acetymethylcarbinol not produced. Starch not hydrolyzed. Small amount of acid and gas from glu- cose and maltose. Trace of acid but no gas from sucrose. No acid or gas from lactose. Acid, gas and ethyl alcohol produced in small quantity from wort together with a pronounced parsnip-like odor. Optimum pH 5.0. No growth at pH 4.0. Temperature relations : Optimum 32°C. FAMILY ACHROMOBACTERIACEAE 439 Good growth at 18° C. Thermal death point 54° C for five minutes. Aerobic, facultative. Distinctive character : Extreme pleo- niorphism in media of neutral or slightly alkaline reaction. Source: Isolated from brewers' yeast. Habitat: The common short rod bac- terium of brewers' yeast. 25. Flavobacterium breve (Frankland and Frankland) Bergey et al. (Kurzer Canalbacillus, Mori, Ztschr. f. Hyg., 4, 1888, 53; Bacillus h-evis G. and P. Frankland, Microorganisms in Water, 1894, 429; not Bacillus brevis Migula, Syst. d. Bakt., 2, 1900, 583; Bacterium breve Chester, Man. Determ. Bact., 1901, 172; Bergey et al.. Manual, 1st ed., 1923, 116.) From Latin brevis, short. Rods: 0.8 to 1.0 by 2.5 microns, show- ing polar staining. Xon-motile. Gram- negative. Gelatin colonies: Minute, pale yellow, compact growth in 2 to 3 weeks. Gelatin stab : Thin, yellowish growth on surface. Beaded growth in stab. No liquefaction. Agar slant : Yellowisli growth in 2 to 3 days. Broth : Turbid with white sediment. ■ Blood serum: Growth of light gray color in 2 to 3 days. Litmus milk: Unchanged. Potato : Xo growth. Aerobic, facultative. Optimum temperature 35°C. Habitat : Water. 26. Flavobacterium solare (Lehmann and Neimiann) Bergey et al. {Bac- terium solare Lelmiann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 258; Bergey et al.. Manual, 1st ed., 1923, 116.) From Latin Solaris, solar. Rods: 0.3 to 0.4 by 1.0 micron, occur- ring singly, in pairs and in chains. Non- motile. Gram-negative. Gelatin colonies : Circular, yellow, glis- tening, translucent. Projecting rays. Gelatin stab : Yellow, arborescent growth in stab. No liquefaction. Agar slant : Pale yellow, raised, arbor- escent. Broth : Clear. Litmus milk: Unchanged. Potato: Soft, yellowish-brown streak. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 30°C. Distinctive character : Resembles Flavobacterium arborescens in type of growth. Source : WiirzlDurg tap water. Gib- bons (Contrib. to Canadian Biol, and Fish., 8, No. 22, 1934, 279) reports this species as occurring in the slime of a skate (Raja erinacea) and of a hake (Urophycis tenuis). Habitat : Fresh and salt water. 27. Flavobacterium flavotenue Schrire. (Trans. Royal Soc. South Africa, 17, 1928, 45.) From Latin flavus, tawny yellow and tenuis, slender. Probably intended to mean, yellow and slender. Small rods: Non-motile. Gram-nega- tive. Agar colonies: Small, circular, lemon yellow, raised, entire. Agar slant : Filiform, lemon yellow. Broth: Turbid. Litmus milk: Unchanged. Potato: Moist, yellow streak. Indole not formed. Nitrites are produced from nitrates. Acid from glucose, galactose and xylose. Pathogenic to white mice and guinea pigs. Aerobic, facultative. Optimum temperature 20°C. Source : Isolated fro m a mold-like growth in a frog {Xenopus laevis). Habitat: Unknown. Appendix : Some of the following spe- cies were described before Gram and flagella stains had been perfected. 440 MANUAL OF DETERMINATIVE BACTERIOLOGY Hence it is impossible to identify them definitely as belonging in Flavohacterium. Comparative study is needed in some cases before other species listed here can be placed in their proper place in the genus. Flavohacterium acidificiun Steinhaus. (Jour. Bact., ^2, 1941, 772.) From the intestine of the grasshopper {Cono- cephalus fasciatus De G.), the Colorado potato beetle {Leptinotarsa decemlineata Say), several unidentified lady beetle larvae, and the white cabh»age butterfly (Pier is rapae L.). Flavohacterium anlenniforme (Ravenel) Bergey et al. (Bacillus antenniformis Ravenel, Memoirs Nat. Acad. Sci., 8, 1896, 25; Bacterium antenniformis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 91; Bergey et al.. Manual, 1st ed., 1923, 104.) From soil. See Manual, 5th ed., 1939. 531 for a description of this organism. Flavohacterium ■ aurantiacum (Frank- land and Frankland) Bergey et al. (Bacillus aurantiacus G. and P. Frank - land, Zeitschr. f. Hyg., 6, 1889, 390; Bacterium aurantiacus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 109; Bergey et al., Manual, 1st ed., 1923, 107; Chromobacterium aurantiacum Top- ley and Wilson, Princ. Bact. and Immun., 7 , 1 93 1 , 405 . ) From wate r . See Man ual , 5th ed., 1939, 533 for a de.scription of this organism. Flavohacterium aurantinum (Hanuner) Bergey et al. (Bacillus aurantinus Ham- mer, Research Bull. No. 20, Iowa Exp. Sta., 1915, 149; Bergey et al.. Manual, 1st ed., 1923, 107.) From milk. See Manual, 5th ed., 1939, 541 for a descrip- tion of this organism. Flavohacterium aurescens (Ravenel) Bergey et al. (Bacillus aurescens Rave- nel, Memoirs Nat. Acad. Sci., 8, 1896, 8; not Bacillus aurescens Frankland and Frankland, Philo. Trans. Roy. Soc. London, B, 1878, 271; Bacterium aures- cens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta.. 9. 1897. 105; Bergey et al.. Manual, 1st ed.. 1923, 1)2.) Fro a joil. Gibbons (Contrib. Canadian Biol, and Fish., 8, No. 24, 1934, 307) found this species in fillets of haddock (Melano- grammus aeglefinus) . See Manual, 4th ed., 1934, 142 for a description of this organism. Flavohacterium brunneum (Copeland) Bergey et al. (Bacillus hrunneus Cope- land, Rept. Filtration Commission, Pitts- burgh, 1899, 348; Bergey et al.. Manual, 1st ed., 1923, 112.) From water. See Manual, 5th ed., 1939, 541 for a descrip- tion of this organism. This may be Bacillus hrunneus Schroeter, but not Bacillus hrunneus Eisenberg. The latter forms spores. Flavohacterium huccalis (Chester) Bergey et al. (Bacillus g, Vignal, Arch, d. phys. norm, et path., Ser. 3, 8, 1886, 365 ; Bacillus huccalis minutus Sternberg, Manual of Bact., 1893, 643; Bacterixim. huccalis mitiulus Chester, Ann. Rept. Del. Col. Agr. Expt. Sta., .9, 1897, 108; Bacterium vignali Migula, Syst. d. Bakt., 2, 1900, 443; Bacterium bucallis (sic) Chester, Man. Determ. Bact., 1901, 167; not Bacterium huccale Migula, Syst. d. Bakt., 2, 1900, 445; Bacillus vignalis Nepveux, These Fac. Pharm., Paris and Nancy, 1920, 112; Bergey etal.. Manual, 1st ed., 1923, 113.) From saliva. See Manual, 5th ed., 1939, 541 for a description of this organism. Flavohacterium hutyri Bergey et al. (Bacillus aromaticus hutyri Severin, Cent. f. Bakt., II Abt., 11, 1903, 264; Bergey et al., Manual, 1st ed., 1923, 106.) From sour cream . Produces an agreeable odor. See Manual, 5th ed., 1939, 534 for a description of this organism. Flavohacterium chlorum Steinhaus. (Jour. Bact., 42, 1941, 772.) From the intestine of the nine-spotted lady beetle (Coccinella novemnotala Habst.). Flavohacterium denitrificans (Lehmann and Neumann) Bergey et al. (Bacillus denitrificans I, Burri and Stutzer, Cent, f. Bakt., II Abt., /, 1895, 360; Bacterium denitrificans I, Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 77; Bac- terium denitrificans Lehmann and Neu- FAMILY ACHROMOBACTERIACEAE 441 mann, Bakt. Diag., 2 Aufl., 2, 1899, 273; Pscudomonas stutzeri Miu;ul:i, Syst. d. Bakt., 2, 1900, 929; Bacillus dcnitrificans Chester, Man. Determ. Bact., 1901, 224; Bergey et al., Manual, 1st ed., 1923, 109; Chromohacterium denitrificans Topley and Wilson, Princ. Bact. and Immun., 1, 1931, 405.) From horse manure. See Manual, 5th ed., 1939, 534 for a descrip- tion of this organism. Flavohacterium desidiusuni (Wright ) Bergej- et al. {Bacillus decidiusus (sic) Wright, Memoirs Nat. Acad. Sci., 7, 1895, 443; not Bacillus desiduosus McBeth, Soil Sci., /, 1916, 450; Bacterium desidio- sus and Bacterium decidiosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 107 and 133; Flavobacterium decidu- osum Bergey et al., Manual, 1st ed., 1923, 114.) From water. Gibbons (Contrib. Canadian Biol, and Fish.. S, No. 24, 1934, 308) found this species in fillets of had- dock (Melanogrammus aeglcfinus) . See Manual, 5th ed., 1939, 514 for a descrip- tion of this organism. Flabobacterium flarescens (Pohl) Bergey et al. (Bacillus flavescens Pohl, Cent. f. Bakt., 11, 1892, 144; Bergey et al.. Manual, 1st ed., 1923, 107.) From water. See Manual, 5th ed., 1939, 535 for a description of this organism. Flavobacterium flavum (Fuhrmann) Bergey et al. {Bacillus flavus Fuhr- mann, Cent. f. Bakt., II Abt., 19, 1907, 117; Manual, 1st ed., 1923, 101.) From beer. See Manual, 4th ed., 1934, 141 for a description of this organism. Flavobacterium gelatinum Sanborn. (Jour. Bact., 19, 1930, 376.) From sea water. Flavobacterium halmephilum Elazari- Volcani. (Studies on the microflora of the Dead Sea, Thesis, Hebrew Univ., Jerusalem, 1940, VIII and 85.) From the Dead Sea. A yellow halophilic species. Flavobacterium halophihim Bergey et al. (Culture No. 30 of Baranik-Pikow- sky, Cent. f. Bakt., II Abt., 70, 1927, 373; Bergey et al., Manual, 3rd ed., 1930, 147.) From sea water. See Manual, 5th ed., 1939, 540 for a description of this organism. Flavobacterium lacunatum (Wright) Bergey et al. {Bacillus lacunatus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 435; Bacterium lacunatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 110; Bergey et al.. Manual, 1st ed., 1923, 117.) From water. See Manual, 5th ed., 1939, 552 for a description of this organism. Flavobacterium matzoonii (Chester) Bergey et al. (Species No. 46 of Conn, Storrs Agr. Exper. Sta., 7th Ann. Rept. for 1894, 1895, 80; Bacillus mata- zooni (sic) Chester, Man. Determ. Bact., 1901, 236; Bergey et al., Manual, 1st ed., 1923, 107.) From matzoon, a fermented milk from Armenia. See Man- ual, 5th ed., 1939, 536 for a description of this organism. Flavobacterium ovale (Wright) Bergey et al. {Bacillus ovalis Wright, Memoirs Nat. Acad. Sci., 7, 1895, 435; Bacterium ovalis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 111; not Bac- terium ovale Migula, Syst. d. Bakt., 2, 1900, 458; Bergey et al.. Manual, 1st ed., 1923, 117.) From water. See Manual, 5th ed., 1939. 551 for a description of this organism. Flavobacterium plicatum (Zimmer- mann) Bergej' et al. (Bacillus plicatus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, Chemnitz, /, 1890, 54; not Bacillus plicatus Frankland and Frank- land, Phil. Trans. Roy. Soc. London, 178, B, 1887, 273; Bergey et al.. Manual, 1st ed., 1923, 105.) From water. Gram- negative. Non-motile. See Manual, 5th ed., 1939, .532 for a description of this organism. See p. 684. Flavobacterium pruneaeum Sanborn. (Jour. Bact., 19. 19.30, 376.) From sea water. Flavobacterium radiaturn (Zimmer- mann) Bergey et al. (Bacillus radiatus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, Chemnitz, 1, 1890, 58; Bacillus radiatus aquatilis Frankland and Frankland, Microorg. in Water, 442 MANUAL OF DETERMINATIVE BACTERIOLOGY London, 1894, 458; Bergey et al.. Manual, 1st ed., 1923, 104.) From water. See Manual, 5th ed., 1939, 531 for a descrip- tion of this organism. Gram-variable. Slight motility of shorter rods. Flavobacterium schirokikhii (H. Jen- sen) Bergey et al. (Salpeter zerstoren- den Bacillus, Schirokikh, Cent. f. Bakt., II Abt., 2, 1896, 205; Bacterium schiro- kikhi H. Jensen, ibid., 4, 1898, 409; Bacillus denitrificans Chester, Man. Determ. Bact., 1901, 274; Bergey et al.. Manual, 1st ed., 1923, 100). From horse manure. See Manual, 5th ed., 1939, 527 for a description of this organism. Flavobacteriiirn stolanatum (Adametz and Wichmann) Bergey et al. {Bacillus stolonaius Adametz and Wichmann, Mitt. Oest. Versuchsstat. f. Brauerei u. Malz., Wien, Heft 1, 1888, 884; Bacterium stolon- atus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 76; Bergey et al., Man- ual, 1st ed., 1923, 106.) See Manual, 5th ed., 1939, 535 for a description of this organism. From water. Flavobacteriwn tremelloides (Tils) Bergey etal. {Bacillus tremelloides Tils, Ztschr. f. Hyg., 9, 1890, 292; Bacterium tremelloides Chester, Ann. Kept. Del. Col. Agr. E.xp. Sta., 9, 1897, 105; Bergey et al.. Manual, 1st ed., 1923, 105.) From river water at Freiburg. Forms a yellow, slimy milk. See Manual, 5th ed., 1939, 532 for a description of this organism. Flavobacteriwn {Halobacterium) maris- mortui Elazari-Volcani. (Studies on the Microflora of the Dead Sea, Thesis, Hebrew Univ., Jerusalem, 1940, V and 48.) From the Dead Sea. This species and Flavobacterium {Halobacterium) halobium and Flavobacterium {Halo- bacterium) trapanicum are placed in a new subgenus of Flavobacterium named Halobacterium. All produce red pig- ment. The flagellation of these species was not determined. They may be polar flagellate, see Pseudomonas salin- aria and P. cuiirubra. Flavobacterium {Halobacterium) halo- bium (Fetter) Elazari-Volcani. (Mi- crobe du rouge de morue, Le Dantec, Compt. rend. Soc. Biol., Paris, 58, 1902, 136; Bacillus halobius ruber Klebahn, Mitteil. a. d. Inst, f . allg. Bot. Hamburg, 4, 1919, 47; Bacterium halobium Fetter, Over rood en andere bacterien van gesouten visch, Diss., Utrecht, 1932; Elazari-Volcani, Studies on the Micro- flora of the Dead Sea, Thesis, Hebrew Univ., Jerusalem, 1940, V and 59.) From reddened salted codfish. Flavobacterium {Halobacterium) tra- 'panicum (Fetter) Elazari-Volcani. {Bac- terium trapanicum Fetter, Over rood en andere bacterien van gezouten visch, Diss., Utrecht, 1932; Elazari-Volcani, Studies on the Microflora of the Dead Sea, Thesis, Hebrew Univ. Jerusalem, 1940, V and 59.) From the Dead Sea. FAMILY ENTEROBACTERIACEAE 443 FAMILY X. ENTEROBACTERIACEAE RAHN. (Cent. f. Bakt., II Abt., 96, 1937, 280.) Gram-negative straight rods. Motile with peritrichous flagella, or non-motile. Grow well on artificial media. All species attack glucose forming acid, or acid and visible gas (H2 present). Characteristically nitrites are produced from nitrates (exceptions in Erwinia only). Antigenic composition is best described as a mosaic which results in serological interrelationships among the several genera, even extend- ing to other families. Many animal parasites, and some plant parasites causing blights and soft rots. Frequently occur as saprophytes causing decomposition of plant mate- rials containing carbohydrates. Note : Early attempts to develop a satisfactory basis for the recognition of species among the coliform-dysentery-typhoid group of bacteria are reviewed by Winslow, Kligler and Rothberg (Jour. Bact., 4, 1919, 429). These were largely based on differences in motility, production of indole, ability to liquefy gelatin, and, more particularly, differences in the ability to ferment carbohydrates, especially such com- pounds as glucose, lactose, sucrose, dulcitol and salicin. The more recent attempts to express differences in species of coliform bacteria by means of the IMViC reac- tion are reviewed by Parr (Amer. Jour. Public Health, 26, 1936, 39; Bact. Rev., 3, 1939, 1), this cryptic symbol indicating the indole test, methyl red acid deter- mination, acetylmethylcarbinol production (Voges-Proskauer reaction) and the util- ization of salts of citric acid. Stuart, Griffin and Baker (Jour. Bact., 36, 1938, 391) and Griffin and Stuart (Jour. Bact., ^O, 1940, 83) have applied these tests plus cellobiose fermentation to a study of a long series of cultures. Capsulated types of coliform bacteria are still placed in this edition of the Manual in a separate genus, Klebsiella, although there is some question about the separation of these from the species in Escherichia and Aerohacter. Meanwhile, the Kauffmann and White Antigenic Schema has been successfully applied to the recognition of serological groups and types among salmonellas and related organisms. The groupings recognized are outlined in the Salmonella Sub- committee Reports submitted to the 2nd and 3rd Congresses of Microbiology (Jour. Hyg., 34, 1934, 333 and Proc. 3rd Internat. Cong, for Microbiology, 1940, 832). The successful use of antigenic structure in this field has stimulated a study of the use of H and O antigens as a means of classifying the coliform group (Stuart, Baker, Zimmerman, Brown and Stone, Jour. Bact., 40, 1940, 101) but this method of classi- fying the species of coliform bacteria has not proved particularly helpful as yet. During this same period there has been an increasing appreciation of the closeness of the relationship between certain common chromogenic bacteria (Serratia) and the coliform bacteria (Breed and Breed, Cent. f. Bakt., II Abt., 71, 1927, 435). INIoreover, the close relationship between bacteria producing soft rots of living vege- table and other plant tissue (now included in Erwinia) and the coliform bacteria has become more evident in recent studies (Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 435). Many intermediate types are found in rotting vegetable materials, these rotting types having the ability to attack protopectin (Burkey, Iowa State Coll. Jour Sci., 3, 1928, 57) but not to cause soft rots of living plant tissue. Borman, Stuart and Wheeler (Jour. Bact., 48, 1944, 351) have proposed a rearrange- ment of the species in thg family Enterobacteriaceae which combines many forms that have previously been regarded as separate species, or even as belonging in sepa- rate genera. Only the future can determine which of all of these views best expresses the relationships of the bacteria belonging in the Family Enterobacteriaceae. — The Editors. 444 MANUAL OF DETERMINATIVE BACTERIOLOGY Key to the tribes of family Enterobacteriaceae. I. Ferment lactose with the formation of acid and visible gas within 24 hours at 37°C or within 48 hours at 25° to 30°C. Some transitional forms produce acid and gas from lactose slowly. Tribe I. Eschericheae, p. 444. II. Plant parasites. Ferment lactose with formation of acid, or acid and visible gas. Usually attack middle lamellar substance in plant tissues, causing soft rots. Tribe II. Erwineae, p. 463. III. Ordinarily chromogenic producing a pink, red or orange-red pigment. Occa- sionall}^ non-pigmented. Ferment glucose and lactose with formation of acid, or acid and visible gas. Tribe III. Serrateae, p. 479. IV. Lactose not fermented within 30 days either at 37°C or at 25° to 30°C. Urea decomposed within 48 hours. Tribe IV. Proteae, p. 486. V. Lactose rarely fermented within 30 days either at 37°C or at 25° to 30°C. Urea not decomposed within 48 hours. Tribe V. Salmonelleae, p. 492. TRIBE I. ESCHERICHEAE BERGEY, BREED AND MURRAY. (Preprint, Manual, 5th ed., October, 1938, vi.) Ferment glucose and lactose with the formation of acid and visible gas within 24 hours at 37°C, or within 48 hours at 25° to 30°C. Some forms produce acid and gas from lactose slowly (occasionally not at all). Do not liquefy gelatin except slowly in Aerobacter cloacae. Key to the genera of tribe Eschericheae.* I. Acetylmethylcarbinol not produced. Methyl red test positive. Salts of citric acid may or may not be used as a sole source of carbon. Genus I. Escherichia, p. 444. II. Acetylmethylcarbinol produced. Methyl red test negative. Salts of citric acid used as sole source of carbon. Genus II. Aerobacter, p. 453. III. Acetylmethylcarbinol may or may not be produced. Methyl red test variable. Salts of citric acid may or may not be used as sole source of carbon. Gas not as abundant as in previous genera. Capsulated forms from respiratory, intes- tinal and genito-urinary regions. Genus III. Klebsiella, p. 457. Genus I . Escherichia Castellani and Chalmers\ . (Castellani and Chalmers, Manual Trop. Med., 3rd ed., 1919, 941; Colibacterium Orla-Jensen, Jour. Bact., 6, 1921, 272; Colobactrum (in part) Borman, Stuart and * Levine (Jour. Bact., /, 1916. 153) was the first to show the inverse correlation be- tween the methyl red and Voges-Proskauer tests and used these characters for the primary separation of the Escherichia coli section and the Aerobacter aerogenes section (Amer. Jour. Public Health, 7, 1917, 784). t Completely revised by Prof. M. W. Yale, New York State Experiment Station, Geneva, New York, Nov., 1938; further revision, July, 1943. ■FAMILY p:xtp:koba(teriaceae 445 Wheeler, Jour. Bact., 4^. 1944. 357.) Named for Theodor Escherich, who first iso- lated the type species. Short rods fermenting glucose and lactose with acid and gas production. Acetyl- methylcarbinol is not produced. Methyl red test positive. Carbon dioxide and hydrogen produced in approximately equal volumes from glucose. Generally not able to utilize uric acid as a sole source of nitrogen. Found in feces and is occasionally pathogenic to man (colitis, cystitis, etc.). It is, however, also widely distributed in nature . The type species is Escherichia coli (Migula) Castellani and Chalmers. Key to the species of genus Escherichia. I. Citric acid and salts of citric acid not utilized as sole source of carbon. A. Hydrogen sulfide not produced. 1. Escherichia coli. II. Citric acid and salts of citric acid utilized as sole source of carbon. A. Hydrogen sulfide produced. 2. Escherichia freundii, B. Hydrogen sulfide not produced. 3. Escherichia intermedium. 1. Escherichia coli (Migula) Castellani and Chalmers. {Bacterium coli com- viune Escherich, Die Darmbakterien des Neugeborenen und Sauglings, 1885; Ba- cillus escherichii Trevisan, I generi e le specie delle Batteriacee, 1889, 15 ; Bacillus coli communis Sternberg, Manual of Bacteriology, 1893, 439; Bacillus coli Migula, in Engler and Prantl, Xattir- lichen Pflanzenfam., /, la, 1895, 27; Bacterium coli Lehmann and Xeumann, Bakt. Diag., 1 Aufi., 2, 1896, 224; Bacillus coli verus Durham, Jour. Exp. Med., 5, 1900, 371; Bacillus coli communis verus Durham, ibid., 353; Aerobacter coli Beijerinck, Cent. f. Bakt., II Abt., 6, 1900, 193; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 941; Bacillus coli-communis Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 483; Bacterium coli-communis Holland, Jour. Bact., 5, 1920, 217; Colobactrum coli Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 358.) From Latin colon, the large intestine. Note: Weldin (Iowa State Jour. Sci., 1, 1927, 121) considers the following identical with the above : Bacillus cavi- cida Fliigge, Die Mikroorganismen, 1886, 268 or more probably Brieger, Berlin. klin. Wochnschr., 1884, No. 14; Bacillus C, Booker, Trans. Ninth Internat. Med. Congress, 3, 1887, 598; Bacillus schafferi von Freudenreich, Landw. Jahrb. d. Schweiz, 4, 1890, 17; Bacterium cavicida Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 130; Bacterium schafferi Chester, ibid., 74; Bacillus mustelae septicus Matzuschita, Bakt. Diag., 1902; Bacillus communis Jackson, Jour. Inf. Dis., 8, 1911, 241 ; not Bacillus communis Migula, Syst. d. Bakt., 2, 1900, 725; Escherichia cavicida Castellani and Chal- mers, Manual of Trop. Med., 3rd ed., 1919, 942; Escherichia schaefferi Bergey et al., :VIanual, 1st ed., 1923, 196. Oesterle (Cent. f. Bakt., I Abt., Orig., 134, 1935, 115) has described a yellow strain Bacterium coli flavum, Parr (Proc. Soc. Exp. Biol, and Med., 35, 1937, 563) a golden-brown strain Bacterium aurescens (not Bacterium aurescens Migula, Syst. d. Bakt., ^, 1900, 466), and Tittsler (Jour. Bact., 33, 1937, 450) reddish-orange strains which are regarded as pigmented variants of Escherichia coli. Rods : Usually 0.5 by 1.0 to 3.0 microns, varying from almost coccoid forms to long rods, occurring singly, in pairs and short chains. Motile or non-motile. Motile 446 MANUAL OF DETERMINATIVE BACTERIOLOGY strains have peritrichous flagella. Not usually capsulated. Non-spore-forming. Gram-negative. Gelatin colonies: Opaque, moist, gray- ish-white, entire. Gelatin stab: Grayish-white, spread- ing, undulate. No liquefaction. Agar colonies: Usually white, some- times yellowish-white, rarely yellow, yellow-brown, golden-brown, reddish- orange or red; entire to undulate, moist, homogeneous. Atypical forms occur fre- quently. Agar slant: Usually white, sometimes yellowish-white, rarely yellow, yellow- brown, golden-brown, reddish-orange or red growth; moist, glistening, spreading. Broth: Turbid, with heavy grayish sediment. No pellicle. Litmus milk: Rapid acid formation with development of gas, usually coagula- tion, curd may or may not be broken up, no peptonization of the curd. Litmus may or may not be reduced. Potato: Abundant, grayish to yellow- ish-brown, spreading. Indole usually formed. Nitrites produced from nitrates. Blood agar plates: Different strains vary widely in their action, some being hemolytic (Buchgraber and Hilk6, Cent, f. Bakt., I Abt., Grig., 1S3, 1935, 449). Heat resistance : Usually destroyed in 30 minutes at 60°C, but certain heat- resistant strains may withstand this exposure (Ayers and Johnson, Jour. Agr. Res., 3, 1914, 401; Stark and Patterson, Jour. Dairy Sci., 19, 1936, 495). Antigenic structure : An antigenically heterogeneous species. Methyl red test positive (Clark and Lubs, Jour. Inf. Dis., 17, 1915, 160); Voges-Proskauer test negative (Durham, Jour. Exp. Med., 5, 1901, 373); inverse correlation between methyl red and Voges-Proskauer tests (Levine, Jour. Bact., 1, 1916, 153). Citric acid and salts of citric acid not utilized as sole source of carbon (Koser, Jour. Bact., 8, 1923, 493). Uric acid not utilized as sole source of nitrogen (Koser, Jour. Inf. Dis., 28, 1918, 377) ; uracil utilized as sole source of nitrogen (Mitchell and Levine, Jour. Bact., 35, 1938, 19). Gas ratio : Approximately equal vol- umes of carbon dioxide and hydrogen, ratio 1:1, produced from glucose (Harden and Walpole, Proc. Roy. Soc, Ser. B, 77, 1905, 399; Rogers, Clark and Davis, Jour. Inf. Dis., 14, 1914, 411). Catalase produced. No H2S produced in peptone iron agar (Levine, Epstein and Vaughn, Amer. Jour. Public Health, 24, 1934, 505; Titts- ler and Sandholzer, Amer. Jour. Public Health, 27, 1937, 1240). More sensitive indicators give positive tests for H2S (Hunter and Weiss, Jour. Bact., 35, 1938, 20). Trimethyleneglycol not produced from glycerol by anaerobic fermentation (Braak, Onderzoekingen over Vergisting van Glycerine, Thesis, Delft, 1928, 166; Werkman and Gillen , Jour. Bact., 23, 1932, 167). Acid and gas from glucose, fructose, galactose, lactose, maltose, arabinose, xylose, rhamnose and mannitol. Sucrose, raffinose, salicin, esculin, dulcitol and glycerol may or may not be fermented. Variable fermentation of sucrose and salicin (Sherman and Wing, Jour. Bact., 33, 1937, 315; Tregoning and Poe, Jour. Bact., 34, 1937, 473) . Inulin, pectin and adonitol rarely fermented. Dextrin, starch, glycogen and inositol not fer- mented. Cellobiose (Jones and Wise, Jour. Bact., 11, 1926, 359) and a-methyl- glucoside (Koser and Saunders, Jour. Bact., 24, 1932, 267) not fermented. Certain strains produce variants which ferment lactose slowly or not at all (Ren- nebaum, Jour. Bact., 30, 1935, 625). Some strains of slow-lactose-fermenters appear to be intermediate between the coliform and paratyphoid groups (Sandi- ford, Jour. Path, and Bact., 4I) 1935, 77). See Twort (Proc. Royal Soc. Lon- don, 79, 1907, 329) for utilization of unusual glucosides; Dozois et al. (Jour. Bact., 30, 1935, 189 and 32, 1936, 499) FAMILY EXTEROBACTERIACEAE 447 for utilization of certain sugar alcohols and their anhydrides; Poe and Klemme (Jour. Biol. Chem., 109, 1935, 43) for utilization of rare sugars. See Winslow, Kligler and Rothberg (Jour. Bact., 4, 1919, 429) for review of literature relative to classification. Fecal odor produced. Aerobic, facultative. Growth requirements: Good growth on ordinary laboratory media. Optimum growth temperature 30° to 37°C. Growth takes place at 10°C and at 45°C. Gas produced from glucose at 45° to 46°C. Eijkmann test positive (Eijkmann, Cent, f. Bakt., I Abt., Orig., 37, 1904, 74; Perry and Hajna, Jour. Bact., 26, 1933, 419). Source : From feces of infants. Habitat : Normal inhabitant of the intestine of man and all vertebrates. Widely distributed in nature. Fre- quently causes infections of the genito- urinary tract. Invades the circulation in agonal stages of diseases. la. Escherichia coli var. acidilactici (Topley and Wilson) Yale. (Milchsaure bacterium, Hueppe, Mit. d. kais. Gesund., 2, 1884, 340; Bacillus acidi lactici Zopf, Die Spaltpilze, 1885, 87; not Bacterium acidi lactici Zopf, Die Spaltpilze, 1884, 60; Bacillus acidi lactici I and // Grotenfelt, Fortschr. d. Med., 7, 1889, 121; possibly also Bac- terium acidi lactici I and // Grotenfelt, ibid., 123; Bacterium acidi lactici Migula, in Engler and Pi-antl, Natiirlichen Pflan- zenfamilien, 1, la, 1895, 25; not Bac- terium acidi lactici Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 357; not Bacterium B, Peters, Bot. Zei- tung, 47, 1889, 422) ; possibly Bacterium grotenfeldtii Migula, Syst. d. Bakt., 2, 1900, 408, a synonym of Bacterium acidi lactici I Grotenfelt; Bacillus acidi- lactici Jackson, Jour. Inf. Dis., 8, 1911, 241 ; possibly Bacillus lacticus IVIac6, Traits pratique de bact., 1913, 452; not Bacillus lacticus Kruse, in Fliigge, Die Mikroorganismen, 2, 1896, 356; Bacterium duodenale Ford, Studies from Victoria Hospital, Montreal, /, 1903, 17 (according to Perkins, Jour. Inf. Dis., 37, 1925, 247) ; Encapsulatus acidi lactici Castellani and Chalmers, Manual of Trop. Med., 1919, 934; Bacillus lactici-acidi Holland, Jour. Bact., 5, 1920, 218; Bacterium acidi- lactici Holland, ibid.; (Encapsulata) Bacillus duodenale Perkins, Jour. Inf. Dis., 37, 1925, 247; Escherichia acidi- lactici Bergey et al., Manual, 1st ed., 1923, 199; Bacterium coli var. acidi lactici Topley and Wilson, Princip. Bact. and Immun., 1, 1931, 446; Yale, in Manual, 5th ed., 1939, 393.) Identification : Includes strains of Escherichia coli which do not attack either sucrose or salicin. It is generally thought that Hueppe's cultures were contaminated with a spore-former. Source : From milk. lb. Escherichia coli var. neapolitana (Topley and Wilson) Yale. (Neapeler Bacterien, Emmerich, Deut. med. Wchnschr., 10, 1884, 299 ; Bacillus neapol- itanus Fliigge, Die Mikroorganismen, 1886, 270; Bacterium neapolitanus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 138; Escherichia neapolitana Castel- lani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 942; Bacterium coli var. nea- politanum Topley and Wilson, Princip. Bact. and Immun., 1, 1931, 446; Yale, in Manual, 5th ed., 1939, 393.) Identification : Includes strains of Escherichia coli which ferment sucrose and salicin. Source : From cholera patients or cad- avers, originally thought to be the cause of cholera. Ic. Escherichia coli var. communior (Topley and Wilson) Yale. {Bacillus coZicommwnior Durham, Jour. Exp. Med., 5, 1900, 353; Bacillus communior Ford, Studies from Victoria Hosp., Montreal, 1, 1903, 17; Bacterium communior Jack- son, Jour. Inf. Dis., 8, 1911, 241 ; Bacillus coli -communior Holland, Jour. Bact., 6, 1920, 217; Bacterium coli-communior 448 MANUAL OF DETERMIXATIVK BACTERIOLOGY Holland, idem; Escherichia communior Bergey et al., Manual, 1st ed., 1923, 200; Bacterium coli var. communior Topley and Wilson, Princip. Bact. and Immun., /, 1931, 446; Yale, in Manual, 5th ed., 1939, 393.) Yale (Cornell Vet., 23, 1933, 306) re- gards Bacterium astheniae Dawson (15th Ann. Rpt., Bur. Anim. Ind., U.S.D.A., 1898, 329; Bacillus astheniae Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 487; Escherichia astheniae Bergey et al., Manual, 1st ed., 1923, 205) as a synonym of Escherichia communior. Identification : Includes strains of Escherichia coli which ferment sucrose but not salicin. Levine (Iowa Eng. Exp. Sta. Bui. 62, 1921, 38) recognizes a strain which ferments salicin. 2. Escherichia freundii (Braak) Yale. {Bacterium, freundii Braak, Onderzoekin- gen over vergisting van glycerine. Thesis, Delft, 1928, 140; Citrobacter freundii Workman and Gillen, Jour. Bact., 23, 1932, 176; Yale, in Manual, 5th ed., 1939, 394; Colobactrum freundii Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 358.) Named for A. Freund, who first observed that tri- methyleneglycol was a product of fer- mentation (1881). Minkewitsch (Ztschr. f. Hyg., Ill, 1930, 180) proposed the name Bacterium coli citrovorum for the intermediates but this name is not acceptable since it is a trinomial . Werkman and Gillen (Jour. Bact., 23, 1932, 177) emended the description of Bacterium freundii, and created the genus Citrobacter. The following species re- named by Werkman and Gillen are regarded as identical with Escherichia freundii: Citrobacter album, Citrobacter decolor ans, Citrobacter diver sum and Citrobacter anindolicum . Tittsler and Sandholzer (Jour. Bact., 29, 1935, 349) and Carpenter and Fulton (Amer. Jour. Pub. Health, 27, 1937, 822) suggest that the intermediates which give a positive methyl red and a negative Voges-Proskauer test be allocated to the genus Escherichia. Other strains are apparently more nearly related to the genus Aerobacter than to the genus Escherichia since they produce acetyl- methylcarbinol. Barritt (Jour. Path. and Bact., 42, 1936, 441 ; 44, 1937, 679) has shown that some of the intermediates form traces of acetylmethylcarbinol which can be detected by the a-naphthol test, but not by the standard Voges- Proskauer test as described in the Manual of Methods for the Pure Culture Study of Bacteria (Soc. Amer. Bact., 1937, 17). Rods : Short rods with rounded ends, occurring singly, in pairs and short chains. Motile or non-motile. Gram- negative. Gelatin stab : Liquefaction by 4 out of 15 cultures (Werkman and Gillen, Jour. Bact., 23, 1932, 177). No liquefaction by any strains (Tittsler and Sandholzer, Jour. Bact., 29, 1935, 353; Carpenter and Fulton, Amer. Jour. Pub. Health, 27, 1937,822). Agar slant: Smooth, gray, shining, filiform and butyrous growth. Litmus milk: Acid in 2 days; coagula- tion may or may not take place ; no pep- tonization. Potato: Abundant, yellowish-white growth. Indole may or may not be formed (Werkman and Gillen, loc. cit. ; Tittsler and Sandholzer, loc. cit.). Nitrites produced from nitrates. Methyl red test positive. Voges- Proskauer test negative (Koser, Jour. Bact., 9, 1924, 59). Some strains give a positive methyl red and a positive Voges- Proskauer test (Parr, Jour. Bact., 36, 1938, 1). Citric acid utilized as sole source of carbon; uric acid not utilized as the sole source of nitrogen (Koser, loc. cit.; Werk- man and Gillen, loc. cit., 167). Catalase produced. Hydrogen sulfide produced in proteose peptone, ferric citrate agar (Levine, FAMILY ENTEROBACTERIACEAE 449 Epstein and Vaughn, Amer. Jour. Pub. Health, 2J^, 1934, 505; Tittsler and Sand- holzer, Amer. Jour. Pub. Health, 27, 1937, 1240). Trimethyleneglycol produced from glycerol by anaerobic fermentation (Braak, loc. cit., 146; Werkman and Gillen, loc. cit., 167). Acid and gas from glucose, fructose, galactose, arabinose, xylose, rafRnose, lactose, maltose, mannose, rhamnose, trehalose, glycerol, mannitol and sorbitol. Sucrose, salicin, dulcitol, adonitol and inositol may or may not be fermented. Cellobiose usually fermented while a- methyl-glucoside may or may not be fermented (Tittsler and Sandholzer, loc. cit.; Carpenter and Fulton, loc. cit.). No acid or gas from amygdalin, dextrin, erythritol, glycogen, inulin or melezitose. Aerobic, facultative. Growth requirements : Good growth on ordinary laboratory media. Optimum growth temperature 30° to 37°C. Gas not produced in Eijkman test when car- ried out at 45° to 46°C (Levine, Epstein and Vaughn, loc. cit.). No gas at 44°C (Wilson, Med. Res. Council, London, Special Rept. Ser. 206, 1935, 165). Habitat : Normally found in soil and water and to a varying degree in the intestinal canal of man and animals. Widely distributed in nature. 3. Escherichia intermedium (Werkman and Gillen) Vaughn and Levine. {Citro- bacter intermedium Werkman and Gillen, Jour. Bact., 23, 1932, 178; Vaughn and Levine, Jour. Bact., U, 1942, 498.) Citrohacter glycologenes W^erkman and Gillen {loc. cit.) is also regarded as a synonym of Escherichia intermedium. Vaughn and Levine (loc. cit.) give a new description of Escherichia inter- medium based on a study of 27 cultures. Rods: Short rods with rounded ends. Occurring singly, in pairs and short chains in young nutrient agar or broth cultures. Actively motile with peritrichous flagella or non-motile. Gram -negative. Gelatin stab : No liquefaction after 60 days at 20°C. Agar slant : Smooth to wrinkled surface, grayish-white, abundant, raised and butyrous growth. Nutrient broth : Turbid with slight ring at surface. Litmus milk : Acid, sometimes coagula- tion and reduction, no proteolysis. Potato: Growth abundant, white to ivory color. Levine's eosine -methylene blue agar: Well-isolated colonies vary from 1 to 4 mm in size. No confluence of neighbor- ing colonies. Colonies are slightly to moderately raised with surfaces varying from flat to convex and usually smooth and glistening but sometimes dull, rough and granular. By transmitted light two types of colonies have been observed : (1 ) Colonies having almost the same appearance throughout but with a distinctly lighter center, the color being similar to the medium. (2) Colonies having a dark brownish central area which diifuses out to a lighter margin. By reflected light three types of col- onies have been observed: (1) Dark, button-like, concentrically ringed col- onies possessing a strong, greenish- metallic sheen so characteristic for Escherichia coli. (2) Colonies with dark, purplish, wine-colored centers surrounded by a light pink zone. Some colonies are concentrically ringed. (3) Pink colonies with no suggestion of sheen but sometimes concentrically ringed. Indole may or may not be formed. Nitrites produced from nitrates. Fermentation of glucose : The end prod- ucts characteristic for the genus Escheri- chia are formed. Carbon dioxide and hydrogen gases are formed in approxi- mately equimolar proportions (gas ratio 1:1) besides significant quantities of ethyl alcohol, and acetic, lactic and succinic acids with only traces of formic acid. Acetylmethylcarbinol and 2-3 450 MANUAL OF DETERMINATIVE BACTERIOLOGY butylene glycol liave not been found (Voges-Proskauer test negative). Salts of citric acid are utilized as a sole source of carbon. Catalase produced. Hydrogen sulfide not detected in proteose peptone ferric-citrate agar. Acid or acid and gas produced from xylose, arabinose, rhamnose, glucose, fructose, mannose, galactose, lactose, maltose, trehalose and mannitol. No acid or gas from melezitose, amygdalin and erythritol. Sucrose, raffinose, cel- lobiose, a-methyl-glucoside, adonitol, dulcitol, glycerol, inositol, sorbitol, starch, aesculin, salicin and sodium malonate may or may not be fermented. Aerobic, facultative. Temperature requirements : Growth at 10°C and at 45° to 46°C. Optimum growth temperature 30° to 37 °C. Gas not produced in Eijkman tests, although some cultures show growth at 45° to 46°C. Salt tolerance : Most cultures ferment glucose in the presence of sodium chloride in a concentration of 6.0 to 7.0 per cent. A few cultures tolerate 8.0 per cent sodium chloride. pH range: Optimum about pH 7.0. Growth occurs at pH 5.0 to pH 8.0. Habitat : Normally found to a varying degree in soil, water and in the intestinal canal of man and animals. Widely dis- tributed in nature. Appendix: The following described species have been placed in Escherichia or may belong here : Bacillus alcalescens Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 37; also see Jour. Med. Res., 6, 1901, 211; not Bacillus alkalescens Andrews, Lancet, 194, 1918, 560; Escherichia alcalescens Bergey et al., Manual, 1st ed., 1923, 202.) From feces. Bacillus asiaticus Castellani. (Cas- tellani. Cent. f. Bakt., I Abt., Orig., 65, 1912, 262; not Bacillus asiaticus Sak- haroff, Ann. Inst. Past., 8, 1893, 550; Salmonella asiaticus Castellani and Chal- mers, Manual of Trop. Med., 3rd ed., 1919, 940; Proteus asiaticus Bergey et al.. Manual, 1st ed., 1923, 211; Bacterium asialicum Weldin and Levine, Abst. Bact., 1, 1923, 13.) From feces. Fer- ments lactose slowly or not at all. Bacillus asiaticus mobilis Castellani. (Valerie 21, Boycott, Jour. Hyg., 6, 1906, 33; Castellani, Ann. di Med., Nav. e Colon., 11, 1916, 453; Salmonella asiaticus mobilis Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 940; Bac- terium valeriei Weldin and Levine, Abst. Bact., 7, 1923, 13; Proteus valeriei Bergey et al.. Manual, 1st ed., 1923, 211.) From feces. A motile variety which Alves (Jour. Path, and Bact., U, 1937, 485) found to be identical with Bacillus asiaticus. Bacillus chylogenes Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 62; also see Jour. Med. Res., 6, 1901, 219.) From feces. Bacillus coli immobilis Kruse . (Kruse , in Fliigge, Die Mikroorganismen, 3 Aufl., 3, 1896, 339; Bacterium coli immobilis Chester, Del. Agr. Exp. Sta., 9th Ann. Rept., 1897, 128; Enteroides eniericus Castellani, Jour. Hyg., 7, 1907, 1 ; Bacillus schafferi MacConkey, Jour. Hyg., 9, 1909, 86; not Bacillus schafferi von Freuden- reich, Landwirtschl. Jahrb. den Schweiz, 4, 1890, 17; Bacillus entericus Castellani and Chalmers, Manual of Trop. Med., 1st ed., 1910, 990; not Bacillus entericus Ford, Studies from Royal Victoria Hosp., Montreal, /, (5), 1903, 40; Escherichia schaefferi Bergey et al.. Manual, 1st ed., 1923, 196; Bacterium coli var. immobilis Winslow et al.. Jour. Bact., 4, 1919, 486; Bacterium schafferi Weldin and Levine, Abst. Bact., 7, 1923, 13; not Bacterium schafferi Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74; Escherichia enterica Weldin, Iowa State Coll. Jour. Sci., 1, 1927, 134.) From feces. These were all described as non-motile variants of Escherischia coli (see Weldin, loc. cit.). Bacillus coli mutabilis Neisser. FAMILY ENTEROBACTERIACEAE 451 (Neisser, Cent. f. Bakt., I Abt., Ref. (Supp.), 38, 1906, 98; Bacterium colt mutabile Massiiii, Arch. f. Hyg., 61, 1907, 250 ; Escherichia coli mutabilis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 943; Escherichia coli-mutabile Deere et al.. Jour. Bact., 31, 1936, 625.) From feces. An unstable variant closely related to Escherichia coli characterized by irregular lactose fermentation. When cultured on lactose indicator agar, it appears not to ferment lactose. After some days, lactose-fermenting papillae appear growing on or out of the original colonies. Subcultures from these sec- ondary colonies give typical lactose fermentation but subculture from the primary colony, avoiding contact with the papillae, gives delayed fermentation of lactose and when again plated will pro- duce non-fermenting colonies on which fermenting papillae later appear. Bacillus gastricus Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 58; also see Jour. Med. Res., 6, 1901, 213; Escherichia gastrica Bergey et al., Manual, 1st ed., 1923, 203.) From feces. Bacillus gruenthali Morgan. (Das gruenthaler Bacterium, Fischer, Z.tschr. f . Hyg., 39, 1902, 447; Morgan, Brit. Med. Jour., 1, 1905, 1257; Bacillus acidi lactici var. gruenthali Levine, Jour. Bact., 3, 1918, 270; Bacterium acidilactici var. gruenthali Winslow, Kligler and Roth- berg, Jour. Bact., 4, 1919, 486; Escherichia gruenthali Castellani and Chalmers, Manual of Trop. Med., 1919, 942; Bac- ierium gruenthali Weldin and Levine, Abst. Bact., 3, 1923, 13.) From feces. Bacillus iliacus Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 61 ; also see Jour. Med. Res., 6, 1901, 213; Escherichia iliacus Bergey et al.. Manual, 1st ed., 1923, 203; Proteus iliacus Bergey et al., IManual, 4th ed., 1934, 363.) From feces. Bacillus infrequens Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 42; also see Jour. Med. Res., 6, 1901, 219.) From feces. Bacillus jejunales Ford. (Ford, Stud- ies from the Royal Victoria Hosp., Mon- treal, /, (5), 1903, 66; also see Jour. Med. Res., 6, 1901, 219.) From feces. Bacillus leporis Migula. {Bacillus leporis lethalis Sternberg, Manual of Bacteriology, 1893,4^53; Bacteriu7n leporis lethalis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 97; Migula, Syst. Bakt., 2, 1900, 651; Eberthella leporis Bergey et al., Manual, 1st ed., 1923, 229; Escherichia leporis Bergey et al., Manual, 2nd ed., 1925, 221.) From feces. Bacillus para-gruenthali Castellani. (Castellani, 1914, quoted from Castellani and Chalmers, Ann. Past. Inst., 34, 1920, 614; Escherichia paragruenthali Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 942; Bacterium coli var. paragruenthali Weldin and Levine, Abst. Bact., 3, 1923, 13.) From feces. Weldin and Levine (Iowa State Coll. Jour. Sci., 1, 1926, 132) regard this species as identical with Bacillus gruenthali Morgan. Bacillus plebeius Ford. (Ford, Stud- ies from the Royal Victoria Hosp., Mon- treal, 1, (5), 1903, 41 ; also see Jour. Med. Res., 6, 1901, 213; Escherichia plebeia Bergey et al., Manual, 1st ed., 1923, 203.) From feces. Bacillus subalcalescens Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, /, (5), 1903, 37; also see Jour. Med. Res., 6, 1901, 217.) From feces. Bacillus subgastricus Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 58; also see Jour. Med. Res., 6, 1901, 219.) From feces. Bacillus vekanda Castellani. (Castel- lani, Jour. Trop. Med. and Hyg., 20, 1917, 181 ; Enteroides vekanda Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 941; Bacterium vekanda Weldin and Levine, Abst. Bact., 7, 1923, 13; Escherichia vekanda Bergey et al., Manual, 1st ed., 1923, 197.) From feces. Bacillus vesiculiformans Henrici. 452 MANUAL OF DETERMINATIVE BACTERIOLOGY (Henrici, Arb. Bakt. Inst. Hochsch. Karlsruhe, 1, 1894, 25; Escherichia vesicu- liformans Bergey et al., Manual, 2nd ed., 1925, 222.) From cheese. Bacterium chijmogenes Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 63; also see Jour. Med. Res., 6, 1901, 219.) From feces. Bacterium coli alcaligenes Chiari and Loffler. (Cent. f. Bakt., I Abt., Orig., 96, 1925, 95.) From feces. Bacterium coli aniyidolicum. Lembke. (Lembke, Arch. f. Hyg., 26, 1896, 299; Bacillus anindolicuin Chester, Man. Determ. Bact., 1901, 207; Escherichia anindolica Bergey et al., Manual, 3rd ed., 1930, 325.) From feces. Bacterium coli imperfectum Roelcke. (Cent. f. Bakt., I Abt., Orig., 145, 1939, 109.) From feces. Lactose not fer- mented. Bacterium Jormicum Omelianski. (Omelianski, Cent. f. Bakt., II Abt., 11, 1904, 184; Achromobacter Jormicum Bergey et al., Manual, 1st ed., 1923, 144; Escherichia formica Bergey et al.. Man- ual, 2nd ed., 1925, 220.) From soil. Bacterium galactophilum Ford. (Ford, Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 39; also see Jour. Med. Res., 6, 1901, 217; Escherichia galactophila Bergey et al.. Manual, 1st ed., 1923, 202.) From feces. Bacterium succinicum Sakaguchi and Tada. (Cent. f. Bakt., II Abt., 101, 1940, 341.) From cheese. Bacterium uromutabile Koch. (Cent, f. Bakt., I Abt., Orig., 1S3, 1935, 209.) From genito-urinary infections. A non- lactose-fermenting variety that devel- oped the ability to ferment lactose slowly. Bacterium, vesiculosum Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 37; Bacillus vesiculosus MacConkey, Jour. Hyg., 9, 1909, 86; Escherichia vesiculosa Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 942.) From cheese. Escherichia alba Schrire. (Trans. Royal Soc. So. Africa, 17, 1928, 43.) From feces. Escherichia brasiliensis Mello. (Sao Paulo Medico, Anno 10, 2, 1937, 11.) From feces. Escherichia colofoetida (Castellani) Hauduroy et al. {Bacillus colofoetidus Castellani, Jour. Trop. Med. and Hyg., 1930, 134; Hauduroy et al., Diet. d. Bact. Path., 1937, 226.) From feces. Escherichia coloides (Castellani) Cas- tellani and Chalmers. {Bacillus coloides var. A and Bacillus coloides var. B, Castellani ; Castellani and Chalmers, Manual of Trop. Med., 1919, 942 and 946.) From feces. Escherichia colotropicalis (Castellani) Castellani and Chalmers. {Bacillus colo- tropicalis Castellani, 1907; see Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 942 and 946.) From feces. Escherichia ellingeri (Metalnikov and Chorine) Bergey et al. {Coccobacillus ellingeri Metalnikov and Chorine, Ann. Inst. Past., 42, 1928, 1635; Bergey et al., Manual, 3rd ed., 1930, 330.) Causes fatal infection in insects as Pyrausta nubilalis Hiibn. (European corn borer) and Gallerin mellonella L. (bee moth). See Manual, 5th ed., 1939, 606 for a de- scription of thig' species. Escherichia khartoumensis (Chalmers and Macdonald) Hauduroy et al. {Ba- cillus khartoumensis Chalmers and Mac- donald, 1915; see Castellani and Chal- mers, Manual of Trop. Med., 3rd ed., 1919, 948; Enieroides khartoumensis Cas- tellani and Chalmers, ibid., 941 ; Haudu- roy et al.. Diet. d. Bact. Path., 1939, 230.) From feces. Escherichia metacoli (Castellani) Cas- tellani and Chalmers. {Bacillus metacoli Castellani, 1915; see Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 942 and 948.) From feces. Escherichia metacoloides (Castellani) Castellani and Chalmers. {Bacillus meta- coloides Castellani; see Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919. 942 and 950.) From feces. FAMILY EXTEROBACTERIACEAE 453 Escherichia paradoxa (Toumanoff) Hauduroy et al. {Colibacillus paradoxus Toumanoff, Bull. Soc. Centr. de Med. V^t^r., 80, 1927, 367; Hauduroy et al., Diet. d. Bact. Path., 1937, 231.) From feces. Escherichia paraenterica (Castellani) Hauduroy et al. (Bacillus paraentericus Castellani, Manual of Trop. Med., 1st ed., 1910, 991; Enteroides paraenterica Castellani and Chalmers, ibid., 3rd ed., 1919, 941; Hauduroy et al.. Diet. d. Bact. Path., 1937, 231.) From feces. Escherichia pauloensis Mello. (Ass. Paulista de Medicina, 11, 1937, 73.) From feces. Escherichia pseudocoli (Castellani) Castellani and Chalmers. {Bacillus pseudo-coli Castellani, iVIanual of Trop. Med., 1st ed., 1910, 990; Castellani and Chalmers, Manual Trop. Med., 3rd ed., 1919, 942.) From feces. Escherichia pseudo-coliformis (Castel- lani) Hauduroy et al. {Bacillus pseudo- coliformis Castellani, 1917; see Castellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 952; Hauduroy et al., Diet. d. Bact. Path., 1937, 233.) From feces. Escherichia pseiidocoloides (Castellani) Castellani and Chalmers. {Bacillus pseudocoloides Castellani, 1916; see Cas- tellani and Chalmers, Manual of Trop. Med., 3rd ed., 1919, 954; ibid., 942; Bacterium pseudo-coloides Weldin and Levine, Abst. Bact., 7, 1923, 13.) From feces. Escherichia pseudocoscoroba Castellani and Chalmers. {Bacillus coscoroba Mac- Conkey, Jour. Hyg., 6, 1906, 570; not Bacillus coscoroba Tr^trop, Ann. Inst. Past., 14, 1900, 224; Bacterium coscorobae Bergey and Deehan, Jour. Med. Res., 19, 1908, 182; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 942; Bacillus communior var. coscoroba Wins- low, Kligler and Rothberg, Jour. Bact., 4, 1919, 486; Escherichia coscoroba Weldin, Iowa State Coll. Jour. Sci., 1, 1926, 139.) From feces and sewage. This organism described by MacConkey is quite different from the organism described bj' Tr^trop (see Pasteurella appendix). Escherichia pseudodysenteriae Bergey et al. {Bacterium pseudodysenteriae Kruse, Deutsche Med. Wchnschr., 27, 1901, 386; Bergey et al., Manual, 1st ed., 1923, 198.) From feces of normal persons and of dysentery patients. Genus II. Aerobacter Beijerinck.* (Beijerinck, Cent. f. Bakt., II Abt., 6, 1900, 193; Aerogenesbacterium Orla-Jensen, Jour. Bact., 6, 1921, 272; Colobactrum (in part) Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 357.) From Latin, air or gas, and rod. Short rods, fermenting glucose and lactose with acid and gas production. Methyl red test negative; Voges-Proskauer test positive. Form two or more times as much carbon dioxide as hydrogen from glucose; trimethyleneglycol not produced from glycerol by anaerobic fermentation; citric acid and salts of citric acid utilized as sole source of carbon. Grow readily on ordinary media. Facultative anaerobes. Widely distributed in nature. The type species is Aerobacter aerog'enes (Kruse) Beijerinck. Note : Kligler (Jour. Inf. Dis., 15, 1914, 187) found the fermentation of glycerol to be inversely correlated with gelatin liquefaction and considered the former the more reliable due to occasional loss of gelatin liquefying ability. This was confirmed by Levine (Amer. Jour. Pub. Health, 7, 1917, 784) who reports that the two characters do not correlate perfectly. Griffin and Stuart (Jour. Bact., 40, 1940, 93ff.) findasimilar correlation of characters but feel that because these characters do not correlate per- fectly, it would be better to combine the two species into a single species. * Completely revised by Prof. M. W. Yale, New York State Experiment Station, Geneva, New York, Nov., 1938; further revision, July, 1943. 454 MANUAL OF DETERMINATIVE BACTERIOLOGY Key to the species of genus Aerobacter. I. Glycerol fermented with acid and gas. A. Gelatin not liquefied (rarely liquefied). 1. Aerobacter aerogenes. II. Glycerol fermented with no visible gas. A. Gelatin liquefied. 2. Aerobacter cloacae. 1. Aerobacter aerogenes (Kruse) Bei- jerinck. {Bacterium lactis aerogenes Es- cherich, Fortschr. d. Med., 3, 1885, 515; Bacterium lactis Baginsky, Ztschr. f. phys. Chem., 12, 1888,437; not Bacterium lactis Lister, Quart. Jour. Micro. Sci., 13, 1873, 380; Bacterium aceticum Bagin- sky, ibid.; Bacillus lactantium Trevisan, I generi e le specie delle Batteriacee, 1889, 15; Bacillus lactis aerogenes Stern- berg, Manual of Bacteriology, 1893, 447; Bacillus aerogenes Kruse, in Fliigge, Die Mikroorganismen, 2, 1896, 340; not Bacillus aerogenes Miller, Deutsche med. Wchnschr., 12, 1886, 119; Bacterium aerogenes Chester, Del. Agr. Exp. Sta., 9th Ann. Rept., 1897, 53; not Bacterium aerogenes Miller, loc. cit.; Beijerinck, Arch. n6erl. d. sci. exact, et nat., 4, 1900, 1 ; Encapsulatus lactis-aerogenes Castel- lani and Chalmers, Manual of Trop. Med., 1919, 934; {Encapsulata) Bacillus aero- genes Perkins, Jour. Inf. Dis., 37, 1925, 254; Colobaclrum aerogenes Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 358.) From Latin, gas-producing. Rods: 0.5 to 0.8 by 1.0 to 2.0 microns, occurring singly. Frequently capsu- lated. (A variety showing a transverse arrangement of the capsule has been named Aerobacter iranscapsulatus by Thompson, Jour. Bact., 28, 1934, 41.) Usually non-motile. Gram -negative. Gelatin colonies: Thick, porcelain- white, opaque, moist, smooth, entire. Gelatin stab: Thick, spreading, white, opaque surface growth. No liquefaction. Agar colonies: Thick, white, raised, moist, smooth, entire. More convex than colonies of Escherichia coli and often mucoid. Agar slant: Abundant, thick, white, moist, glistening, spreading growth. Broth : Turbid, with pellicle and abun- dant sediment. Litmus milk: Acid with coagulation. No peptonization. Potato: Thick, yellowish-white to yellowish-brown, spreading with nodular outgrowths over the surface. Indole may or may not be formed (Ford, Studies from the Royal Victoria Hospital, Montreal, 1, 1901-1903, 16; Bardsley, Jour. Hyg. (Eng.), 34, 1934, 38; Wilson, Med. Res. Council, London, Spec. Rept. Ser. 206, 1935, 161). Nitrites produced from nitrates. Methyl red test negative (Clark and Lubs, Jour. Inf. Dis., 17, 1915, 160); Voges-Proskauer test positive (Durham, Jour. Exp. Med., 5, 1901, 373); inverse correlation between methyl red and Voges-Proskauer tests (Levine, Jour. Bact., 1, 1916, 153). Citric acid and salts of citric acid utilized as sole source of carbon (Koser, Jour. Bact., 8, 1923, 493). Uric acid utilized as sole source of nitrogen (Koser, Jour. Inf. Dis., 23, 1918, 377). Gas ratio : Two or more volumes of carbon dioxide to one of hydrogen formed from glucose (Harden and Walpole, Proc. Roy. Soc. Series B, 77, 1905, 399; Rogers, Clark and Davis, Jour. Inf. Dis., 14, 1914, 411). Catalase produced. Hydrogen sulfide not produced in pep- tone iron agar (Levine, Epstein and Vaughn, Amer. Jour. Pub. Health, 24, 1934, 505; Tittsler and Sandholzer, Amer. Jour. Pub. Health, 27, 1937, 1240). More sensitive indicators give positive FAMILY ENTEROBACTERIACEAE 455 tests for hydrogen sulfide (Hunter and Weiss, Jour. Bact., 85, 1938, 20). Trimethyleneglycol not produced from glycerol by anaerobic fermentation (Braak, Onderzoekingen over Vergisting van Glycerine, Thesis, Delft, 1928, 212; Werkman and Gillen, Jour. Bact., 23, 1932, 167). Sodium hippurate hydrolyzed (Hajna and Damon, Amer. Jour. Hyg., 19, 1934, 545). Acid and gas from glucose, galactose, lactose, fructose, arabinose, maltose, rafiinose, cellobiose, salicin, esculin, starch, dextrin, glycerol, mannitol, sor- bitol and inositol, a-methyl-glucoside usually fermented (Koser and Saunders, Jour. Bact., 24, 1932, 267). Sucrose, inulin, dulcitol and adonitol may or may not be fermented. Proto pectin not fer- mented. Variable fermentation of su- crose and mannitol (Sherman and Wing, Jour. Bact., 33, 1937, 315). Aerobic, facultative. Growth requirements : Good growth on ordinary laboratory media. Optimum growth temperature about 30°C. Grows better at temperatures below 30°C than does Escherichia coli. Usually destroj^ed in 30 minutes at 60°C, but certain heat- resistant strains may withstand this exposure (Aj^ers and Johnson, Jour. Agr. Res., 3, 1914, 401 ; Stark and Patterson, Jour. Dairy Sci., 19, 1936, 495). Gas not produced in Eijkmann test when carried out at 45° to 46°C (Eijkmann, Cent. f. Bakt., I Abt., Orig., 37, 1904, 74; Levine, Epstein and Vaughn, Amer. Jour. Pub. Health, 24, 1934, 505). Habitat : Normally found on grains and plants and to a varying degree in the intestinal canal of man and animals. Widely distributed in nature. 2. Aerobacter cloacae (Jordan) Bergey et al. {Bacillus cloacae Jordan, Rept. Mass. State Bd. of Health, Part II, 1890, 836 ; Bacterium cloacae Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 239; Bacillus lactis cloacae Conn, Esten and Stocking, Storrs Agr. Exp. Sta., Conn., 18th Ann. Rept. for 1906, 180; Cloaca cloacae Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 938; Bergey et al.. Manual, 1st ed., 1923, 207.) From Latin cloaca, sewer. The following are also regarded as identical with Aerobacter cloacae: Aero- bacter liquefaciens Grimes and Hennerty, Sci. Proc. Royal Dublin Society, (N. S.) 20, 1931, 93; not Aerobacter liquefaciens Beijerinck, Cent. f. Bakt., II Abt., 6, 1900, 199 (monotrichous) ; Bacillus levans Wolffin, Arch. f. Hyg., 21, 1894, 279 and Lehmann, Cent. f. Bakt., 15, 1894, 350 (Bacterium levans Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 235; Cloaca levans Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 938; Aerobacter levans Bergey et al., Manual, Isted., 1923, 208). Rods: 0.5 to 1.0 by 1.0 to 2.0 microns, occurring singly. Usually motile posses- sing peritrichous flagella. Not capsu- lated. Gram-negative. Gelatin colonies : Thin, circular, bluish, translucent. Gelatin stab : Slow liquefaction. Liquefying power sometimes lost (Klig- ler, Jour. Inf. Dis., 15, 1914, 199). Agar colonies : Circular, thick, opaque with white center, entire. Agar slant: Porcelain-white, smooth, glistening, spreading growth. Broth: Turbid, with thin pellicle. Litmus milk: Acid, coagulation, gas, slow peptonization. Potato : Growth yellowish, moist, glis- tening. Indole not formed (Levine, Epstein and Vaughn, loc. cit.; Wilson, Med. Res. Council, London, Spec. Rept. Ser. 206, 1935, 161). Nitrites produced from nitrates. Methyl red test negative; Voges-Pros- kauer test positive. Citric acid and salts of citric acid utilized as sole source of carbon (Koser, Jour. Bact., 8, 1923, 493). Uric acid utilized as sole source of 456 MANUAL OF DETERMINATIVE BACTERIOLOGY nitrogen (Koser, Jour. Inf. Dis., 23, 1918, 377). Gas ratio : Glucose fermented with lit least two volumes of carbon dioxide to one of hydrogen (Rogers, Clark and Davis, Jour. Inf. Dis., U, 1914, 411). Catalase produced. Hydrogen sulfide not produced in pep- tone iron agar (Levino, Epstein and Vaughn, Amer. Jour. Pub. Health, 24, 1934, 505). Sodium hippurate not hydrolyzed (Hajna and Damon, Amer. Jour. Hyg., 19, 1934, 545). Acid and gas from glucose, fructose, galactose, arabinose, xylosie, lactose, maltose, raffinose, dextrin, salicin, tre- halose, mannitol, sorbitol, cellobiose and a-methyl-glucoside. Sucrose usually fer- mented. Inulin, esculin, starch, dul- citol, rhamnose and proto pectin not attacked. Glycerol fermented with no visible gas (Kligler, loc. cit., 187; Levine, Amer. Jour. Pub. Health, 7, 1917, 784). Starch rarely fermented (Levine, ibid.). See Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 429 for review of literature. Fecal odor produced. Aerobic, facultative. Growth requirements : Good growth on ordinary laboratory media. Optimum growth temperature 30° to 37°C. Gas not produced in Eijkmann test when carried out at 45° to 46°C (Levine, Ep- stein and Vaughn, loc. cit.). Habitat : Found in human and animal feces, sewage, soil and water. Appendix: The following described species have been placed in Aerobacter or may belong here : Actinobacter polymorphus Duclaux. (Duclaux, Ann. Inst. Nat. Agron., 5, 1882, 110; Bacillus actinobacter Migula, Syst. d. Bakt., 2, 1900, 689.) Causes swelling of cheese. Possibly this was Aerobacter cloacae. Aerobacter chinense Bergey et al. {Bacillus capsidatus chinensis Hamilton, Cent. f. Bakt., II Abt., 4, 1898, 230; Bacterium chinense Migula, Syst. d. Bakt., ^, 1900, 357; Bergey etal., Manual, 1st ed., 1923, 207.) From India ink. Aerobacter decolorans Burkey. (Iowa State Coll. Jour. Sci., 3, 1928, 77.) From rotted potato and hay infusions. Aerobacter diver sum Burkey. (Iowa State Coll. Jour. Sci., 3, 1928, 77.) From soil. Aerobacter Jaeni Burkey. (Iowa State Coll. Jour. Sci., 3, 1928, 77.) From hay infusions. Aerobacter hibernicum. Grimes and Hennerty. (Sci. Proc. Royal Dublin Society, (N.S.) 20, 1931, 92.) From butter. Aerobacter leporis Botta. (Giorn. Bat- teriol. e Immunol., 23, 1939, 217.) From liver abscess in a rabbit. Aerobacter melezitovorum Burkey. (Iowa State Coll. Jour. Sci., 3, 1928, 77.) From soil. Aerobacter pectinovorum Burkey. (Iowa State Coll. Jour. Sci., 3, 1928, 77.) From creek water. Aerobacter oxytocum (Trevisan) Ber- gey et al. {Bacillus oxytocus perniciosus Fliigge, Die Mikroorganismen, 1886, 268; Bacillus oxytocus Trevisan, I generi e le specie delle Batteriacee, 1889, 17; Bac- terium oxytocus perniciosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 139; Bacterium oxytocum Migula, Syst. d. Bakt., 2, 1900, 394; Escherichia oxytocus Castellani and Chalmers, Man- ual of Trop. Med., 3rd ed., 1919, 942; Bergey et al.. Manual, 1st ed., 1923, 206.) From old milk. Aerobacter paraoxijtocum Mello. (Jorn. Dos Clinicos, No. 15, 1937.) From a dental abscess. Bacillus aceris Edson and Carpenter. (Edson and Carpenter, Vermont Agr. Exp. Sta. Bull. 167, 1912, 475; Achrotno- bacter aceris Bergey et al.. Manual, 4th ed., 1934, 218.) From slimy maple sap. See Manual, 5th ed., 1939, 506 for a de- scription of this organism. Identified by Fabian (Ind. and Eng. Chem., 27, 1935, 349) as Aerobacter aerogenes. FAMILY EA'TEROBACTERIACEAE -157 Bacillus aromaticus Paminel. (Pam- mel, Bull. No. 21, Iowa Agr. Exper. Sta., 1893, 792; Pammel and Pammel, Cent. f. Bakt., II Abt., 2, 1896, 633; Bacterium aromaticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., .9, 1897, 100; Flavobac- terium aromaticum Bergey et al., Manual, Isted., 1923, 105.) From cabbage. Used as a starter for cheese making. Acid and gas from glucose and sucrose. See Man- ual, 5th ed., 1939, 533 for a description of this organism. Bacillus guillebeau a, b and c, von Freu- denreich. (Ann. de Micrographie, 2, 1890, 353.) From mastitis milk. Cul- ture a may well have been Aerobacter aerogenes, b appears to have been A. cloacae while c was a mucoid variant (see Sternberg, Man. of Bact., 1893, 725). Bacillus subcloacae Ford. (Studies from the Royal Victoria Hosp., Montreal, /, (5), 1903, 60; also see Ford, Jour. Med. Res., 6, 1901, 213.) From feces. Bacterium liquefaciens Ford. (Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 59; also see Ford, Jour. Med. Res., 6, 1901, 215.) From feces. While Ford regards this species as identical with Bacillus liquefaciens Eisenberg, neither is adequately described and they differ in important characters. The same holds true for Bacillus liquefaciens Fuller and Johnson, Jour. Exp. Med., 4, 1899, 627. Bacterium margaritaceum. Migula. (Perlschnurbacillus, Maschek, Bakteriol. Untersuch. d. Leitmeritz. Trinkwaster, Leitmeritz, 1887; Migula, Syst. d. Bakt., 3, 1900, 422 and 1059.) From water. Possibly identical with Aerobacter aero- genes. Bacterium subliquefaciens Ford. (Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 59; also see Ford, Jour. Med. Res., 6, 1901, 219.) From feces. Bacterium zeae Comes. (Bacterial Disease of Corn, Burrill, 111. Agr. Exp. Sta. Bull. 6, 1889, 164; Comes, Critto- gamia Agraria, /, 1891, 500; Bacillus secalis Ludwig, Lehrbuch der niederen Kryptogamen, 1892, 95; Bacillus zeae Russell, Bacteria in their relation to vegetable tissue, Thesis, Johns Hopkins Univ., Baltimore, 1892, 36.) From corn blight. Moore (Agric. Sci., 8, 1894, 368) identified a culture received from Burrill as Bacillus cloacae Jordan. Burkey (Iowa State College Jour. Sci., 5, 1928, 77) described five species {Aero- bacter indologenes, Aerobacter motorium, Aerobacter mitificans, Aerobacter saiici- novorum and Aerobacter pseudoproteus) which are regarded as varieties of Aero- bacter cloacae. Genus III. Klebsiella Trevisan.* (Trevisan, Atti della accad. Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 105; Calymmatobacterium Aragao and Vianno, Mem. Inst. Oswaldo Cruz, 4, 1912, 222; Encapsulatus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 934.) Named for Edwin Klebs (1834-1913), early German bacteriologist. Short rods, somewhat plump with rounded ends, mostly occurring singly. Encap- sulated in the mucoid phase. Non-motile. Gram -negative. Fermentation reactions are highly variable but usually a number of carbohydrates are fermented. Nitrites are produced from nitrates. Aerobic, growing well on ordinary culture media. En- countered frequently in the respiratory, intestinal and genito-urinary tracts of man, but may be isolated from a variety of animals and materials. The type species is Klebsiella pneumoniae (Schroeter) Trevisan. * Rearranged by Prof. M. W. Yale, New York State Experiment Station, Geneva, New York, Nov., 1938; further revision by Dr. O. B. Chapman, Syracuse Medical College, Syracuse, New York, December, 1945. 458 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Klebsiella pneumoniae (Schroeter) Trevisan. (Pneumoniecoccus, Fried- laender, Arch. f. Path. Anat., 87, 1882, 319; Baclerium pneumonie crouposae Zopf, Die Spaltpilze, 3 Aufl., 1885, 66; Klebsiella crouposa Trevisan, Atti della Accad. Fisio-Medico-Statistica in Mil- ano, Ser. 4, 3, 1885, 105; Hyalococcus pneumoniae Schroeter, in Cohn, Krypto- gamen Flora von Schlesien, 3(1), 1886, 152; Bacillus pneumoniae Fliigge, Die Mikroorganismen, 2 Aufi., 1886, 204; Trevisan, Rend. d. R. Istit. Lombardo, Ser. 2, 20, 1887, 94 ; Klebsiella friedlanderi Trevisan, I generi e le specie delle Bat- teriacee, 1889, 26; Bacillus mucosus capsulatus Paulsen, Mittheil. f. d. Verein Schleswig-Holsteiner Aerzte, 2, 1893, No. 7; Bacterium pneumoniae Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 200; Bacterium pneumonicum Mig- ula, Syst. d. Bakt., 2, 1900, 350; Bacillus friedlanderi Mac^, Traits Pratique de Bact., 4th ed., 1901, 771; Encapsulatus pneumoniae Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 934; Coccobacillus friedlanderi Neveu- Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 20; Proteus pneumoniae Weklin, Iowa State Coll. Jour. Sci., 1, 1926, 149; Bacterium friedlander Weldin, idem; Bacillus niucosus-capsulatus Mason and Beattie, Arch, of Internal Med., 42, 1928, 331.) From Greek, of pneumonia. Rods: 0.3 to 0.5 by 5.0 microns, with rounded ends, often four to five times as long as broad, occurring singly and in pairs. Encapsulated. Non-motile. Gram-negative. Gelatin colonies: Dirty-white, smooth, opaque, entire, slightly raised. Gelatin stab : Dirty-white surface growth. Filiform growth in stab. No liquefaction. Gas bubbles. Agar colonies: White, shiny, convex, smooth, glistening, entire. Agar slant: Slimy, white, somewhat translucent, raised growth. Broth : Turbid, with thick ring or film. Litmus milk: Variable. Potato: Yellowish, slimy, raised growth. Gas is formed. Nitrites produced from nitrates. Indole variable, usually not formed. Fermentation of carbohydrates highly variable. Acid and gas may be formed from glucose, lactose, sucrose, fructose, galactose, maltose, mannitol and inositol. Methyl red test variable. Acetylmethylcarbinol production vari- able. Blood agar: Usually no hemolysis. Utilization of citrate as a sole source of carbon variable. Aerobic, facultative. Optimum temperature 37°C. Common name : Friedliinder's pneumo- bacillus. Source : Originally isolated from spu- tum in pneumonia. Habitat : Associated with infections of the respiratory, intestinal and genito- urinary tracts of man. Encountered in infections of animals and may be isolated from a wide variety of sources. Note : The difficulty experienced in distinguishing members of this genus from those of Escherichia and Aerobacter is recognized. The members of these three genera exist in at least three growth phases, mucoid (capsulated), smooth and rough. Working with the mucoid phase of Klebsiella, Julianelle (Jour. Exp. Med., U, 1926, 113, 683, 735; 52, 1930, 539) de- scribed three serological types. A, B and C on the basis of capsular specific polj^- saccharides. There is evidence that other types exist. The presence of a generic specific somatic antigen pattern has not been definitely accepted. Appendix: The following organisms may be placed in Klebsiella. The evi- dence for differentiating them into dis- tinct species is so meagre that for the present it may be better to consider them as varieties of Klebsiella pneumoniae. Klebsiella adanti Hauduroy et al. FAMILY ENTEROBACTERIACEAE 459 (Diet. d. Bact. Path., 1937, 260.) From a case of pyelocystitis. Klebsiella capsulata (Sternberg) Ber- gey et al. (Kapselbacillus, Pfeiffer, Ztschr. f. Hyg., 6, 1889, 145; Bacillus capsulatus Sternberg, Manual of Bact., 1893, 431 ; Bacterium capsulatum IMigula, Syst. d. Bakt., 2, 1900, MQ ; Encapsulatus pfeifferi Bergey et al., Manual, 1st ed., 1923, 239; Bergey et al., Manual, 2nd ed., 1925, 265.) From purulent exudate from stomach of a guinea pig. Klebsiella crassa Trevisan. {Bacillus sputigenus crassus Kreibohm, Inaug. Diss., Gottingen, 1889; abst. in Cent, f. Bakt., 7, 1890, 313; Bacillus crassus sputigenus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 260; Trevisan, I generi e le specie delle Batteriacee, 1889, 25; Bacterium sputigenes crassus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 88; Bacterium crassum Chester, Man. Determ. Bact., 1901, 151.) From sputum . Klebsiella cuniculi Hauduroy et al. {Bacillus capsulatus pyaemiae cuniculi Koppinayi, Ztschr. f. Tiermed., 11, 1907, 429; Hauduroy et al.. Diet. d. Bact. Path., 1937, 262.) From pleuroperi- earditis in a rabbit. Klebsiella genitalium (Dimock and Edwards) Hauduroy et al. {Encapsula- tus genitalium Dimock and Edwards, Jour. Amer. Veter. Assoc, 70, 1927, 469; Hauduroy et al.. Diet. d. Bact. Path., 1937, 264.) ■ From infections in the gen- ito-urinary organs of mares. Klebsiella granulomatis (Aragao and Vianna) Bergey et al. {Calymmatobac- terium granulomatis Aragao and Vianna, Mem. do Inst. Oswaldo Cruz, Rio de Janeiro, 4, 1912, 211; Encapsulatus in- guinalis Bergey et al., Manual, 1st ed., 1923, 238; Bergey et al., Manual, 2nd ed., 1925, 264.) From granuloma inguinale. Klebsiella ozaenae (Abel) Bergey et al. {Bacillus mucosus ozaenae Abel, Cent. f. Bakt., 13, 1893, 167; Bacillus ozaenae Abel, ibid., 172; not Bacillus ozaenae Migula, Syst. d. Bakt., 2, 1900, 645, 662; ? Bacillus capsulatus mucosus Fasching, Sitzgsber. Wien. Akad., Ill Abt., 100, 1891 {Bacterium capsulatus mucosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 130; Bacterium faschingii Migula, Syst. d. Bakt., 2, 1900, 355; Bacillus capsulatus-mucosus Hol- land, Jour. Bact., 5, 1920, 217; Bacterium mucosum capsulatum Holland, ibid., 221 ; Bacterium mucosum -capsulatum Holland, ibid., 217) ; Bacterium ozaenae Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 204; Bacterium mucosus ozaena Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 138; Encapsulata ozenae Bergey et al.. Manual, 1st ed., 1923, 240; Bergey et al., Manual, 2nd ed., 1925, 266.) From cases of ozena. Klebsiella paralytica Cahn, Wallace and Thomas. (Wallace, Lyell, Thomas, Alvin and Cahn, Proc. Soc. Exp. Biol., 29, 1932, 1908; Cahn, Wallace and Thomas, Science, 76, 1932, 385; Wallace, Cahn and Thomas, Jour. Inf. Dis., 53, 1933, 386; Klebsiella aids Hauduroy et al.. Diet. d. Bact. Path., 1937, 260.) From intestine of tick {Dermacentor albipictus) and thought to be the cause of tick paralysis of moose. Klebsiella rhinoscleromatis Trevisan. (Rhinoscleromabaeillus, v. Friseh, Wien. med. Wehnschr., 1882; Cornil, Progrts Medical, 1883; Trevisan, Rend. d. R. Istit. Lombardo, Ser. 2, 20, 1887, 95; Bacterium rhinoscleromatis Migula, Syst. d. Bakt., 2, 1900, 352; Bacterium nasalis Chester, Man. Determ. Bact., 1901, 134; Bacillus rhinoscleromatis Winslow, Klig- ler and Rothberg, Jour. Bact., 4, 1919, 491.) From cases of rhinoscleroma. 460 MANUAL OF DETERMINATIVE BACTERIOLOGY *Appendix I. Tribe Eschericheae: Borman, Wheeler and Stuart (Jour. Bact., 4S, 1944, 361) place coliform-like bacteria that are slow lactose-fermenters in a separate genus Paracolobactrum as follows: Genus A. Paracolobactrum Borman, Stuart and Wheeler. (Paracolibacille, Widal and Nobecourt, Semaine MM., 17, 1897, 285; Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 361.) Short rods characterized by consistently delayed fermentation of lactose (occasion- ally negative). Glucose is fei"mented with formation of visible gas. Certain forms attack carbohydrates characteristically at 20° to 30°C but not at 37 °C. Antigenic relationships to other genera in the family are common, even with respect to major antigens. The type species is Faracolobactrum aerogenoides Borman, Stuart and Wheeler. Key to the species of genus Paracolobactrum. I. Acetylmethylcarbinol produced. 1. Paracolobactrum aerogenoides. II. Acetylmethylcarbinol not produced. A. Citric acid utilized as a sole source of carbon. 2. Paracolobactrum intermedium. B. Citric acid not utilized as a sole source of carbon. 3. Paracolobactrum coliforme. 1. Paracolobactrum aerogenoides Bor- man, Stuart and Wheeler. (Para-aero- genes, Stuart, Wheeler, Rustigian and Zimmerman, Jour. Bact., 43, 1943, 117; Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 361.) Latinized, resembling aerogenes. Characters as for Aerobacter aerogenes and Aerobacter cloacae except for con- sistently delayed fermentation of lactose. Source: From human gastroenteritis. Habitat: Surface water, .soils, grains, as well as the intestinal tract of animals, including man. 2. Paracolobactrum intermedium Bor- man, Stuart and Wheeler. (Para- freundii, Stuart et al., Jour. Bact., 4S, 1943, 117; Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 361.) From Latin intermedins , intermediate . Characters as for Escherichia freundii and Escherichia intermedium except for consistently delayed fermentation of lactose. Source : From human gastroenteritis. Habitat: Surface water, soil, grains, as well as the intestinal tract of animals, in- cluding man. 3. Paracolobactrum coliforme Borman Stuart and Wheeler. (Para-coli, Stuart et al.. Jour. Bact., 45, 1943, 117; Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 361.) Latinized, resembling coli. Characters as for Escherichia coli except for consistently delayed fermen- tation of lactose. Source : From human gastroenteritis. Habitat: Surface water, soil, grains, as well as intestinal tract of animals, in- cluding man. Note : The following also belong here : Bacterium paracoli Stutzer and Wsorow. (Non-lactose-fermenting Bac- terium coli, Gilbert and Lion, Semaine Med., 13, 1893, 130; Stutzer and Wsorow, Cent. f. Bakt., II Abt., 71, 1927, 115.) From intestines of healthy larvae of a moth {Euxoa segetum). Salmonella para-colon (Day) Hauduroy et al. {Bacillus para-colon Day; see Cas- tellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 264; also Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919,950; Hauduroy et al.. Diet. d. Bact. Path., 1937, 461.) * Prepared by Dr. E. K. Borman, Bureau of Laboratories, State Department of Health, Hartford, Connecticut, July, 1945. FAMILY ENT1:R0BACTERIACEAE 461 *Appendix II. Tribe Eschericheae: Gram-negative, peritrichous to non-motile rods similar to organisms placed in Paracolobactrum , Serratia and Salmonella have recently been described as causing diseases of reptiles, birds and mammals. They may be grouped here although they have been placed in several different genera. 1. Bacterium sauromali Conti and Crowley. (Jour. Bact., 36, 1938, 269.) From a generic name of lizards, Sauromalus. Short rods: 0.2 to 0.5 by 1.0 to 2.0 microns, with rounded ends, occurring in groups. Motile with 4 to 6 peri- trichous flagella. Gram-negative. Gelatin: Infundibuliform liquefaction complete in 3 days at 37°C. Black sedi- ment. Medium browned. Agar slant: Growth abundant, spread- ing, convex, faint yellowish-green, glis- tening, smooth, translucent, butyrous. Decided odor. Medium greened. Nutrient broth: After 1 day at 37°C, moderate turbidity. Ring. Decided odor. Scanty flocculent sediment. Milk: Alkaline and complete pepto- nization in ten days. Indole not formed. Potato: Growth yellowish-green to olive. Blood medium: (Complete alpha hemolysis in 48 hours. Peptone medium: Slight fluorescent greenish -yellow pigmentation. Nitrites produced from nitrates. Ammonia is produced. Acid and gas from glucose, sucrose, maltose, galactose, fructose, salicin and mannitol. Acid but not gas from gly- cerol. No acid from lactose, arabinose, xylose, dextrin, inulin. dulcitol or starch. Hydrogen sulfide produced. Catalase positive. Methyl red test positive. Pathogenic for animals. Temperature relations: Optimum 37°C. Minimum 20°C. Maximum 45°C'. Aerobic. Source: From a tumor-like growth on the chuckawalla (Sauromalus varius). Habitat: Causes tumors in lizards. 2. Serratia anolium Duran-Reynals and Clausen. (Jour. Bact., 33, 1937, 369.) From a generic name of lizards, A7iolis. Rods: 0.2 to 0.4 by 1.0 to 2.0 microns, occurring singly, in pairs, in clusters and palisades. Pleomorphic, other forms be- ing 4 to 5 microns in length, curved, occasionally club-like, or 10 to 15 microns long and surrounded by a capsular mate- rial, or occasionally small and coccus-like. Motile (Duran-Reynals and Clausen) with 1 to 4 peritrichous flagella (Breed). Non-acid-fast. Gram -negative. Gelatin stab: Rapid growth. Lique- faction infundibuliform. After 6 to 10 days a thick soft pellicle and blackish sediment is formed. Agar colonies : After 24 hours at 37°C, isolated colonies are low, convex, margin entire or slightly undulate. Colonies translucent, butyrous, glistening, smooth, 1 .0 to 2.5 mm in diameter. While some colonies retain their smooth char- acter, oihers become larger, striated and wrinkled, showing opaque, radiated folds with irregularly crenated edges and a rougher texture. Penetrating acid smell produced. Agar slant : After 24 hours at 37 °C, abundant, confluent, raised, whitish, butyrous, glistening, wuth entire or undulate edges. Broth: Moderate growth with uniform turbidity. A pellicle is formed which disintegrates forming a ring on the walls of the tube. Sediment. Faint fluores- cent yellowish coloration. No visible gas in glucose broth (Breed) . Peptone water: After 6 to 10 days marked turbidity, medium darkened, blackish sediment formed. Litmus milk: Coagulation and diges- tion. Partial discoloration of the litmus. * Prepared by Prof. Robert S. Geneva, New York, June, 1946. Breed, New York State Experiment Station, 462 MANUAL OF DETERMINATIVE BACTERIOLOGY Potato : Growth abundant, butyrous, glistening, raised, pinkish. Indole not formed. Blood is hemolyzed. Loeffler's serum: Abundant, glistening growth. Liquefaction. No H2S produced. Ammonia is produced. Although Duran-Reynals and Clausen report nitrites not produced from ni- trates, a retest of their cultures by Breed has shown that nitrites are actively produced from nitrates. Acid from glucose, fructose, sucrose, mannitol, maltose, galactose and salicin. Dextrin, lactose, inulin, dulcitol, xylose and arabinose slightly attacked or not at all. Pigment production : Water-soluble pigment produced. Pink coloration best shown on glycerol potato. Reddish col- oration best shown in peptone water with 2 per cent glucose, the yellow coloration in glucose broth and the black coloration in the sediment of liquefied gelatin and peptone water. Some non-pigmented strains. Temperature relations: Grows well at 20°C. Growth more abundant at 37°C. Practically no growth at 10°C. Thermal death point 60°C for 20 minutes. Aerobic. Pathogenicity : Pathogenic for am- phibians, reptiles and to some extent fish. Lesions are produced in the iguanid liz- ards (Anolis equestris and Anolis caroli- nensis), the gekkonid lizards {Tarentola mauritanica and Hemidactylus brookii), the garter snake (Thamnophis butleri) and the brown snake {Storeria dekayi), the musk turtle (Sternothaerus odoratus), the toad {Bufo americanus) , the frog (Rana pipiens) and the catfish (Ameiu- ras melas). When the inoculated animal is kept at 37°C, the disease becomes general and usually is fatal. Non-patho- genic for warm-blooded animals (Clausen and Duran-Reynals, Amer. Jour. Path., 13, 1937, 441). Source : From tumor-like lesions in Cuban lizards (Anolis equestris). Also isolated from iguanid lizards (Basiliscus vittatus) from Mexico by Clausen and Duran-Reynals {loc. cit.). Habitat: The cause of a natural, non- fatal, contagious disease of lizards. 3. Salmonella sp. (Type Arizona). {Salmonella sp., Dar es salaam Type var. from Arizona, Caldwell and Ryerson, Jour. Inf. Dis., 65, 1939, 245; Salmonella arizona Kauffmann, Acta Path, et Micro- biol. Scand., 17, 1940, or 19, 1942; Ari- zona culture, Edwards, Cherry and Bru- ner, Jour. Inf. Dis., 73, 1943, 236; Sal- monella arizona Group, Edwards, Jour. Bact., 49, 1945, 513.) Ferments lactose and liquefies gelatin. Antigenic structure: XXXIII: Z4, Z23, zae: — . Source: Isolated by Caldwell and Ry- erson (loc. cit.) from horned lizards, Gila monsters and chuckawallas. Also pathogenic for guinea pigs and rabbits. Found in snakes by Hinshaw and McNeill (Cornell Vet., 34, 1944, 248). Also reported by Edwards (loc. cit.) from infants. Habitat: Apparently widely distri- buted in lizards, snakes, and warm- blooded animals. FAMILY ENTEROBACTERIACEAE 463 TRIBE II. ERWINEAE WINSLOW ET AL. (Jour. Bact., 5, 1920, 209.) Motile rods which normally require organic nitrogen compounds for growth. Pro- duce acid with or without visible gas from a variety of sugars. In some species, the number of carbon compounds attacked is limited and lactose may not be fermented. May or may not liquefy gelatin. May or may not produce nitrites from nitrates. In- vade the tissues of living plants and produce dry necrosis, galls, wilts and soft rots. In the latter case, a pro topee tinase destroys the middle lamellar substance. There is a single genus. Genus I. Erwinia Winslow et al* (Jour. Bact., 2, 1917, 560.) Named for Erwin F. Smith, pioneer American plant pathologist . Characters as for the tribe. The type species is Erwinia amylovora (Burrill) Winslow et al. Key to the species of genus Erwinia. I. **Pathogens which cause dry necrosis, galls or wilts in plants but not a soft rot (Erwinia sensu stricto). A. Gas not produced in sugar media. 1. Gelatin liquefied. a. Starch not hydrolyzed. b. Nitrites not produced from nitrates. 1. Erwinia amylovora. bb. Nitrites produced from nitrates. 2. Erwinia milletiae. aa. Starch hydrolyzed. b. Nitrites produced from nitrates. 3. Erwinia vitivora. aaa. Action on starch not reported. b. Nitrites produced from nitrates. 4. Erwinia cassavae. 2. Gelatin not liquefied. a. Starch not hydrolyzed. b. Nitrites produced from nitrates. 5. Erwinia salicis. bb. Nitrites not produced from nitrates. 6. Erwinia tracheiphila. * Completely revised by Prof. F. D. Chester, New York, N. Y., December, 1938; further revision by Prof. Walter H. Burkholder, Cornell University, Ithaca, New York, May, 1945. ** The genus Erwinia as defined here is heterogeneous in nature and is composed of at least two distinct groups. The first group constitutes Erwinia proper and does not produce visible gas from sugars. Waldee (Iowa State Coll. Jour. Sci., 19, 1945, 435) in a paper that appeared as this manuscript was ready for the press has sug- gested that the species in this first group be placed in a separate family Erwiniaceae. 464 MANUAL OF DETERMINATIVE BACTERIOLOGY II. fPathogens which normally cause a soft rot in plants (largely belong in the genus Pectobacterium Waldee). A. Gas produced in sugar media. 1. Gelatin liquefied. a. Nitrites produced from nitrates, b. Hydrogen sulfide produced. 7. Erwinia betivora. 8. Envinia carnegieana. bb. Hydrogen sulfide not produced. 9. Erwinia atruseptica. 10. Erwinia carotovora. aa. Nitrites not produced from nitrates. 11. Erwinia erivanensis. 2. Gelatin not liquefied, a. Starch hydrolyzed. aa. Starch not hydrolyzed. 12. Erwinia flavida. 13. Erwinia dissolvens. 14. Ernnnin nimipressuralis . B. Gas not produced in sugar media. 1. Gelatin liquefied. a. Nitrites produced from nitrates, b. Starch hydrolyzed. 15. Erwinia ananas. 16. Erwinia cytolytica. bb. Starch not hydrolyzed. c. Acid from lactose. 17. Erwinia aroideae. 18. Erwinia mangiferae. cc. No acid from lactose. 19. Erwinia citrimaculans. 2. Gelatin not liquefied. 20. Erwinia rhapontici. 3. Very slow gelatin liquefaction. a. Nitrites not produced from nitrates. 21. Erwinia lathyri. C. Gas production not reported. 1. Gelatin liquefied. a. Nitrites produced from nitrates. 22. Erwinia lilii. t The second group of species usually causes soft rots, but includes a few not very typical species. Waldee (loc. cit.) has proposed that the species that cause typical soft rot be placed in a new genus, Pectobacterium, with Pectobacterium carotovorum as the type species. The new genus is retained in the family Enter obacteriaceae. Wal- dee would place the atypical species in other genera, Erwinia dissolvens for example being placed in the genus Aerobacter. As further comparative studies are needed before such changes can be made with confidence, the older arrangement is allowed to stand in this edition of the Manual. FAMILY ENTEROBACTERIACEAE 465 1. Erwinia amylovora (Burrill) Wins- low et al. {Micrococcus amylovorus Bur- rill, Illinois Indust. Univ., 11th Rept., 1882, 142; American Naturalist, 17, 1883, 319; Bacillus amylovorus Trevisan, I generi e le specie delle Batteriacee, 1889, 19; Bacterium amylovorus Chester, Del. Col. Agr. Exp. Sta., 9th Ann. Rept., 1897, 127; Bacterium amylovorum Ches- ter, Ivlanual Determ. Bact., 1901, 176; Winslow et al., Jour. Bact., 5, 1920, 209.) From Latin, starch devouring. Description mainlj^ from Ark, Phj-to- path., 27, 1937, 1. Rods: 0.7 to 1.0 by 0.9 to 1.5 microns, occurring singly, in pairs and sometimes in short chains. Motile with peritrichous flagella. Gram-negative. Gelatin colonies: Circular, whitish, amorphous, entire. Gelatin stab: Slow crateriform lique- faction confined to the upper layer. Agar colonies: Circular, grayish-white, moist, glistening, irregular margins. Broth: Turbid, with a thin granular pellicle. Potato: Growth white, moist, glisten- ing. Medium not softened. No odor. No pigment. Litmus milk: Coagulated after 3 to 4 days to a pasty condition, with a separa- tion of whey. At first acid, becoming alkaline. Litmus reduced. There is a gradual digestion of the casein. Blood serum : Growth similar to that on agar. No liquefaction. Dunham's solution: Rapid growth, but clouding not dense. Indole not produced. Nitrites not produced from nitrates. Most of the strains gave a positive test for ammonia in broth, a few showed onlj' a slight positive test. Acetylmethylcarbinol produced. Growth in synthetic media with (NH4)2HP04 as a source of nitrogen and containing different carbohydrates. Acid without gas from glucose, sucrose, arabinose, mannose, fructose, maltose, cellobiose, raffinose, salicin and amyg- dalin. Xylose, rhamnose, dulcitol and starch not fermented. Acid production from lactose and galactose variable. Utilizes salts of citric, malic, and hip- puric acid. Action on salts of lactic and succinic acids variable. Salts of benzoic, maleic, malonic, oxalic, tartaric and val- eric acid are not utilized. Asparagine fermented with production of alkali. Glycine, valine, isoleucine, glutamic acid, cystine, tyrosine, trypto- phane and urea not fermented. Minimum temperature between 3° and S°C. IMaximum below 37 °C. Optimum pH 6.8. Minimum pH 4.0 to 4.4. Maximmn pH 8.8. Source : From the blossoms, leaves and twigs of the pear and apple. Habitat : Attacks a large number of species in several tribes of the famil}' Rosaceae (Elliott, Manual Bact. Plant Pathogens, 1930, 19). 2. Erwinia milletiae (Ivawkami and Yoshida) Magrou. {Bacillus milletiae Ivawkami and Yoshida, Bot. Mag., Tokyo, 34, 1920, 110; INIagrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 213.) From Milletia, a genus named for A. J. Millett. Rods: 0.4 to 0.6 by 0.9 to 2.5 microns. Motile with peritrichous flagella. Cap- sules. Gram -negative. Gelatin: Liquefaction begins after 8 days. Agar colonies : Circular, flat, smooth, shiny, opaque, waxy yellow. Margins entire. Broth: Turbid. Heavy precipitate. Milk: No coagulation. Clears with alkaline reaction. Conjac : No liquefaction. Nitrites produced from nitrates. Acid but no gas from galactose, fruc- tose, lactose, maltose, sucrose and man- nitol. No acid from glycerol. Starch not hydrolyzed. Growth in 0.2 per cent but not in 0.3 per cent of the following acids in sucrose 466 MANUAL OF DETERMINATIVE BACTERIOLOGY peptone broth: Acetic, citric, oxalic and tartaric. Aerobic. Grows well at 32°C. Thermal death point, 53°C for 10 min. Source : From galls on the Japanese wisteria in various localities in Japan. Habitat : Causes galls on the Japanese wisteria, Milletia floribunda. 3. Erwinia vitivora (Baccarini) du Plessis. {Bacillus vilivorus Baccarini, Bull, della Soc. Bot. Ital., 1894, 235; du Plessis, Dept. Agr. and Forestry Union of S. Africa, Science Bui. 214, 1940, 58.) From Latin, devouring the vine. Note: Macchiati (Bol. della Soc. Bot., 1897, 156) uses the name Bacillus bacca- rinii for Bacillus vilivorus. The de- scription Macchiati gives is not of Erwinia vitivora but is evidently that of a saprophyte occurring with the patho- gen. He conducted no inoculation ex- periments. Migula (System der Bak- terien, 2, 1900, 778) gives Bacillus vili- vorus Bacc. (Malpighia, 6, 1892, 229) which is an incorrect citation and Bacil- lus baccarinii Macch. 1897, as synonyms of Bacillus gmumis Comes 1884. It is impossible to determine what this latter species is. Du Plessis {loc. cit.) does not believe Bacillus gummis is the same as Erwinia vitivora. Rods: 0.74 (0.44 to 1.10) by 1.46 (0.95 to 2.19) microns. Cells sometimes dumbbell-shaped. Motile with peritri- chous flagella. Gram-negative. Capsules present. Gelatin: Liquefaction. Agar colonies : first puiictiform, irregu- larly circular or lenticular, ultimately circular, raised to pulvinate, glistening, spreading, light to orange-yellow. Agar becomes brown. Broth: Turbid in 24 hrs. Whitish to lemon yellow pellicle. Milk: Litmus reduced. Thread-like to spongy curd formed. Yellow whey about curd. Yellow growth on top of plain milk. Medium acid. Uschinsky's solution: Slowly becomes turbid. Pellicle. Sediment whitish- yellow. Nitrites produced from nitrates. Hydrogen sulfide produced. Acid produced from glucose, fructose, xylose, lactose, sucrose, mannitol and salicin. No acid from raffinose or inulin. Starch hydrolyzed. Facultative anaerobe. Temperature relations : Optimum 25°C. Maximum 35° to 40°C. Minimum 5° to 10°C. Optinmm pH 6.0. Minimum 4.2. Source : Du Plessis used 5 isolates from various localities in South Africa. Habitat : Causes a disease of grape vines in South Africa, Italy and France. 4. Erwinia cassavae (Handsford) comb. nov. {Bacterium cassavae Handsford, Ann. Ilept. Dept. Agric. Luanda for 1937, II, 1938, 48.) From cassava, the liost plant. Hods : Motile with a few peritrichous Hagella. No capsules. Gram-negative. Gelatin is slowly liquefied. Agar colonies: Smooth, lens-shaped, edges entire, translucent and of uniform structure. Yellow. Broth: Turbid with a ring. A yellow precipitate in old cultures. Milk becomes alkaline. Not cleared. Nitrates are rapidly reduced to ni- trites. Methyl red test negative. Acetyl- methylcarbinol produced (Dowson, Cent, f. Bakt., II Abt., 100, 1939, 183). Acid but no gas from glucose, sucrose, maltose and glycerol, but not from lac- tose . Facultative anaerobe. Source : From necrotic lesions on cas- sava leaves in Uganda. Habitat : Pathogenic on cassava, Mani- hot sp. 5. Erwinia salicis (Day) Chester. {Bacterium salicis Day, Oxford For. Mem., 3, 1924, 14; Phytomonas salicis FAMILY ENTEUOBACTERIACEAE 467 Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 408; Chester, in Ber- gey et al., Manual, 5th ed., 1939, 406.) From Latin salix, willow; M. L. generic name, Salix. Description from Dowson, Ann. Appl. Biol., 24, 1937, 542. Rods: 0.5 to 0.7 by 0.8 to 2.2 microns, occurring singly or in pairs, rarely in chains, with rounded ends. Motile with 5 to 7 long peritrichous flagella. Gram- negative . Gelatin stab: Beaded growth. No liquefaction. Infusion agar: Colonies appear slowly, circular, with slightly uneven margins, pale brown by transmitted light, pale gray by reflected. Infusion agar slants: Growth thin, nearly transparent. Broth: Moderate, uniform turbidit3^ No pellicle. Litmus milk: No change. Potato: Bright yellow, later fading to pale brown, spreading, abundant, glisten- ing, slimy growth. Indole not formed. Nitrites produced from nitrates (Dow- son). Hydrogen sulfide not produced. Ammonia not produced. Acetylmethylcarbinol produced. Methyl red test negative (Dowson, Cent, f. Bakt., II Abt., 100, 1939, 183). Acid, but no gas, from glucose, galac- tose, mannose, xylose, maltose, sucrose, raffinose, glycerol, mannitol and salicin. No growth in arabinose, fructose, rham- nose, inulin or dextrin. No growth in Cohn's solution. Starch not hydrolyzed. Temperature relations: Optimum 29° to 30°C. Minimum 5° to 10°C. Maxi- mum 33° to 37°C. Thermal death point 50° to 52°C. Aerobic, facultative anaerobic. Source : From the cricket-bat willow {Salix caerulea) and from the white willow {Salix alba). Habitat : Causes a water-mark disease of willow in England. 6. Erwinia tracheiphila (Erw. Smith) Holland. {Bacillus tracheiphilus Erw. Smith, Cent. f. Bakt., II Abt., 1, 1895, 364; Bacterium tracheiphilus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 72; Smith, see Bacteria in Rela- tion to Plant Diseases, 2, 1911, 286; Holland, Jour. Bact., 5, 1920, 215.) From Greek, trachea-loving, i.e., live in filirovascular bundles. Rods: 0.5 to 0.7 by 1.2 to 2.5 microns, with rounded ends, occurring singly and in pairs, more rarely in fours. Motile with peritrichous flagella. Capsulated. Gram-negative. Gelatin colonies: Small, circular, gray- ish-white, smooth, glistening. Show in- ternal striae by transmitted light. Gelatin stab: Surface growth thin, spreading, grayish-white. Slight filiform growth in depth. No liquefaction. Agar colonies: Small, circular, grayish- white, smooth, glistening. Agar slant: Growth gray, smooth, filiform, moist, glistening. Broth: Slight turbidity. No ring or pellicle. Potato : Growth white or color of sub- stratum, smooth, moist, glistening. No action on the starch. Does not soften the middle lamella of potato cells. Litmus milk: Not coagulated. Reac- tion unchanged. Litmus not reduced. Not peptonized. Egg albumen: Not digested. Blood serum: No liquefaction. Cohn's solution: No growth. Uschinsky's solution : Weak growth. Fermi's solution: Weak growth. Indole not formed in Dunham's solu- tion. Nitrites not produced from nitrates. Ammonia production moderate. Cannot utilize asparagine, ammonium lactate or tartarate as sources of nitrogen. Acid without gas from glucose, sucrose and fructose; growth in closed arm. Acid from glycerol. No growth in closed arm with lactose, maltose, dextrin, gly- cerol or mannitol. No acid from lactose. Starch not hydrolyzed. 468 MANUAL OF DETERMINATIVE BACTERIOLOGY Hydrogen sulfide production feeble. Growth in broth with 1 .0 per cent NaCl retarded; inhibited with 2.0 per cent. Very sensitive to acid (phenolphtha- lein). Temperature relations : Optimum 25° to 30°C. Minimum about 8°C. Maxi- mum 34° to 35°C. Thermal death point 43 °C for one hour. Aerobe and facultative anaerobe. Source : From various curcurbits. Habitat : Causes the wilt of cucumber, also affects cantaloupes, muskmelons, pumpkins and squashes. 6a. Bacillus tracheiphilus var. cucumis E. F. Smith. (An Introduction to Bact. Dis. of Plants, 1920, 135.) Smith states that squash is immune to this variety of Erwinia tracheiphila. 7. Erwinia betivora (Takimoto) Ma- grou. {Bacillus belivorus Takimoto, Ann. Phyt. Soc. Japan, 2, 1931, 356; Magrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 200). From Latin, devour- ing the beet. Rod: Short rods, sometimes filaments. Motile with 2 to 6 peritrichous flagella. Gram-negative. Gelatin: Liquefaction. Agar colonies: Circular or amoeboid, homogenous, thin, edges smooth and entire. Broth: Turbid with pellicle. Milk: Acid, coagulated. Nitrites produced from nitrates. Indole produced. Hydrogen sulfide produced. Gas from glucose and sucrose. Facultative anaerobic. Optimum temperature 35°C. Minimum 12°C. Maximum 45°C. Thermal death point 50°C for 10 min. Source : From rot of sugar beets in Korea. Habitat : Pathogenic on roots of beets. Artificial inoculation of carrots, radishes, potato tubers and tomato fruits gave positive results. 8. Erwinia carnegieana Lightle, Stand- ring and Brown. (Phytopath., 3£, 1942, 310.) From the genus Carnegiea. Rods: 1.12 to 1.79 by 1.56 to 2.90 microns. Motile with peritrichous flag- ella. Capsules. Gram-positive (Lightle et al.). Gram-negative; old cultures show Gram-positive granules in cells (Burkholder). Gelatin: Slow liquefaction. Agar colonies: Round, slightly raised, smooth, gray-white, wet-shining, mar- gins entire. Broth: Abundant growth. Uschinsky's solution: Turbid, slight ring and sediment. Milk: Litmus pink to reduced. No curdling. Nitrites are produced from nitrates. Hydrogen sulfide is formed (Burk- holder) . Acid and gas from glucose, galactose, fructose, maltose, sucrose, raffinose, mannitol and salicin. Acid and gas from lactose and xylose and alkali from sodium tartrate (Burkholder). Starch not hydrolyzed (Burkholder). No odor. Aerobic. Thermal death point 59°C. Source : From rotting tissue of the giant cactus (Carnegiea gigantia). Habitat : Pathogenic on the giant cac- tus, but not on carrots. 9. Erwinia atroseptica (van Hall) Jennison. (Bacillus atrosepticus van Hall, Inaug. Diss., Amsterdam, 1902, 134; Jennison, Ann. Missouri Bot. Gard., 10, 1923, 43.) From Latin ater, black and septicus, putrefying. Synonyms: Morse (Jour. Agr. Res., 8, 1917, 79) lists the following synonyms: Bacillus solanisaprus Harrison, Cent. f. Bakt., II Abt., 17, 1906, 34 (Erwinia solanisapra Holland, Jour. Bact., 5, 1920, 222) and Bacillus melanogenes Pethy- bridge and Murphy, Roy. Irish Acad. Proc, 29, B, No. 1, 1911, 31. Paine (Jour. Agr. Sci., 8, pt. 4, 1917, FAMILY ENTEROBACTERIACEAE 469 492) agrees and points out that Bacillus phyiophthorus Appel is very similar to Bacillus melanogenes Pethybridge and Murphy. Jennison (Ann. Missouri Bot. Gard., 10, 1923, 1) concurs and adds Bacillus phTjtophthorus Appel, Ber. d. Deut. Bot. Gesell., 20, 1902, 128 (nomen nudum) and K. Biol. Anst. f. Land. u. Forst. Arb., 3, 1903, 364. (This last reference contains Appel's description which is antedated by van Hall's description of the black leg pathogen.) Stapp (Arb. d. Biol. Reichs. f. Land- u. Forst., 16, 1928, 702) besides the above species adds Bacillus carotovorus Jones but uses the name Bacillus phyiophthorus and states that the species contains 5 serological groups. Description from Jennison {loc. cit.). Rods : 0.6 by 1.5 microns. Motile with a few peritrichous flagella. No capsules. Gram-negative. Gelatin liquefied. Agar colonies: Small, round to some- what irregular and whitish. Surface smooth with a glistening luster. Broth : Turbid after a few days. Ring" and sometimes a light pellicle. Ammonia production feeble to moder- ate (Jennison). Ammonia production absent (Morse, loc. cit.). Milk coagulated and acid. A slow peptonization. Litmus reduced. Indole not formed. Hydrogen sulfide not produced. Nitrites are produced from nitrates. Acid and gas from glucose, galactose, sucrose, lactose, maltose and mannitol. No acid and gas from dextrin and glycerol . Volume of gas is small. Starch not hydrolyzed. Cohn's solution: No growth. Uschinsky's solution: Good growth. Facultative anerobe (Morse, loc. cit.). Optimum temperature 26°C. Maxi- mum 33°C. Minimum below 5°C (Morse). Slight growth with 3 per cent salt. None with 4 per cent salt. Source : From stems of potatoes af- fected with black-leg. Habitat: Causes a black rot on stem and tuber of potatoes and other vege- tables. Note: Smith (Science, 31, 1910, 748) regarded Erwinia solanisapra and Er- winia phytophthora as very closely re- lated. Brooks, Nain and Rhodes (Jour. Path, and Bact., 28, 1925, 203) held that Erwinia phytophthora, Erwinia sola- nisapra and Erwinia carotovora are dis- tinct serologically, although identical in cultural characteristics. Berridge (Ann. Appl. Biol., 13, 1926, 12) claimed from serological tests that Erwinia phytophthora and Erwinia solanisapra are different yet closely related organ- isms. Lacey (Ann. Appl. Biol., 13, 1926, 1) from cultural and serological tests considered Erwinia phytophthora, Er- winia solanisapra and Erwinia caroto- vora distinct species. Stapp (Arb. a. d. Biol. Reichanstalt f. Landw. u. Forst- wirtsch., 16, 1928, 643) from serological tests places Erwinia phytophthora in one serological group and Erwinia carotovora in another. Leach (Phytopath., 20, 1930, 743) found that Erwinia phytoph- thora and Erioinia carotovora were in- distinguishable in cultural and physio- logical characteristics, the most con- sistent difference being the inky black coloration of the tissues infected with the former. Stapp (in Sorauer, Handb. d. Pflan- zenk., 5 Aufl., 2, 1928, 229) states that it is generally believed that the disease caused by Bacillus solanincola Delacroix (Compt. rend. Acad. Sci., Paris, 133, 1901, 417 and 1030) is the same as stem rot of potato (blackleg). 10. Erwinia carotovora (Jones) Hol- land. {Bacillus carotovorus Jones, Cent, f. Bakt., II Abt., 7, 1901, 12; Holland, Jour. Bact., 5, 1920, 222; Bacterium caro- tovorum Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 446; Pectobac- 470 MANUAL OF DETERMINATIVE BACTERIOLOGY terium carotovorum Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 469.) From Latin, carrot destroying. Synonyms : Leach (Minnesota Agr. E.xp. Sta. Tech. Bull. 76, 1931, 18) lists the following as synonyms : Bacillus airosepticus van Hall. (Van Hall, Inaug. Diss., Univ. Amsterdam, 1902, 134; Erwinia atroseptica Jennison, Ann. Missouri Bot. Gard., 10, 1923, 43.) Bacillus phijlophthorus Appel. (Ber. d. deut. Bot. Ges., 20, 1902, 128; Erwinia phytophthora Holland, Jour. Bact., 5, 1920, 222; Bacterium phytophthorum Burgwitz, Phytopath. Bacteria, Lenin- grad, 1935, 141; Pectohacterium phyto- phthorum Waldee, Iowa State Coll. Jour. Sci., 19, 1945,471). Bacillus solanisaprus Harrison. (Har- rison, Cent. f. Bakt., II Abt., 17, 1907, 34; Erivinia solanisapra Holland, Jour. Bact., 5, 1920, 222.) Bacillus melanogenes Pethybridge and Murphy. (Roy. Irish Acad., 29, B, No. 1, 1911, 31.) Bacillus oleraceae Harrison. (Harri- son, Science, 16, 1902, 152; Erivinia olera- ceae Holland, Jour. Bact., 5, 1920, 222.) Bacillus omnivorus van Hall. (Inaug. Diss., Univ. Amsterdam, 1902, 176.) Bacillus apivorus Wormald. (Jour. Sci., 6, 1914, 203.) Elrod (Bot. Gaz., 103, 1941, 270) holds that Erwinia aroideae is a synonym of Erivinia carotovora. The following also have been considered as possible synonyms of Erwinia caro- tovora : Bacillus cepivorus Delacroix. (Dela- croix, Ann. Inst. Nat. Agron., Ser 2, 5, 1905, 368; Bacterium cepivorum Stapp, in Sorauer, Handb. d. Pflanzenkrank- heiten, 5 Aufl., 2, 1928, 49; Aplanobacter cepivorus Elliott, Man. Bact. Plant Path., 1930, 4; Phytomonas cepivora Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 344.) Causes a rot of onion bulbs. Bacillus cypripedii Hori. (Hori, Cent. f. Bakt., II Abt., 31, 1911, 85; Erwinia cxjpripedii Bergey et al . , Manual , 1st ed., 1923, 171.) Bacillus dahliae Her. and Bokuii (Hori and Bokura, Imp. Agr. Expt. Sta. Nishigahara, 38, 1911 , 69 ; Erwinia dahliae Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 205.) Pseudomonas destructans Potter. (Potter, Proc. Univ. Durham Philos. Soc, 1899, 165 and Proc. Roy. Soc, 67, 1900, 449; Bacterium destructans Nakata, Nakajima and Takimoto, Tech. Rept. Korea Ind. Farm, 1922; Phytomonas destructans Bergey et al., Manual, 3rd ed., 1930, 264.) See Paine (Ann. Appl. Biol., 5, 1918, 64) for a discussion of this species. Bacillus hyacinthi septicus Heinz. (Heinz, Cent. f. Bakt., 5, 1889, 539; Bacillus hyacinthi-scpticus Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 449 ; Bacterium hyacinthi septicus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 127; Bacillus hyacinthi Migula, Syst. d. Bakt., 2, 1900, 874; not Bacillus hyacinthi Trevisan, I generi e le specie delle Batteriacec, 1899, 19; Ertcinia hyacinthi septica Alagrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 208.) Rods: Usually 0.7 to 0.8 by 1.5 to 5.0 microns, occasionallj^ in chains. Stain rather slowly with aniline colors, well with LofHer's methylene blue. No cap- sules ob.served. .Vctively motile with peritrichous flagella. Gram-negative. Gelatin stab : Rapid surface liquefac- tion, slower in depth (Jones). Some strains very slow liquefiers. Agar colonies: After 2 days, circular, convex, smooth, grayish-white, moist, glistening. Margins sharp, entire. Agar slant : Growth thin, grayish- white, moist, glistening, butyrous. Me- diuni not discolored. Broth : Turbid, with pellicle and white fiocculent sediment. Slow alkaline pro- duction. Litmus milk: After 4 days, coagu- lated, acid, with separation of whey. Cheesy odor. Litmus reduced. Slightly peptonized. FAM ILY FNTEROBACTERIACEAE 471 Potato: Growth thick, creamy-white; medium softened. Dunham's solution: Feeble persistent turbidity. Blood serum : Growth much as on agar. Not liquefied. Uschinsky's solution : Strong turbidity. Indole production none. Nitrites produced from nitrates. Diastase negative. No H2S produced or only a trace. No ammonia produced. Methyl red positive, Voges-Proskauer negative (Dowson, Cent. f. Bakt., II Abt., 100, 1939, 183). Acid and gas from glucose, lactose, sucrose, fructose, raffinose, mannitol, arabinose, .xylose, salicin and rhamnose. Acid without gas from glycerol and ethyl alcohol. Butyl alcohol, inulin and starch not fermented. Facultative anerobe. Temperature relations : Optimum 25° to 30°C. Minimum 4°C. Maximum 38° to 39°C. Thermal death point 41° to 51 °C. Pathogenesis: Causes a rapid soft rot of roots, rhizomes, fruits and the fleshy stems of a variety of plants. Source : From rotted carrots. Habitat : Causes a soft rot in carrot , cabbage, celery, cucumber, egg-plant, iris, muskmelon, hyacinth, onion, pars- nip, pepper, potato, radish, tomato, turnip, and other plants. 11. Erwinia erivanensis (Kalantarian) Bergey et al. {Bacterium erivanense Kalantarian, Cent. f. Bakt., II Abt., 65, 1925, 298; Bacillus einvanensis Stapp, in Sorauer, Handb. d. Pflanzenkr., 5 Aufl., 2, 1928, 202; Bergey et al.. Manual, 3rd ed., 1930, 239.) Derived from Erivan, a city in Armenia. Whether this organism is to be con- sidered a chromogenic strain or a distinct species is impossible to determine ; there- fore, it occupies its present position tentatively. It cannot be separated from Erwinia carotovora on the basis of chromogenesis since the latter occasion- ally shows a tendency to the formation of a faint yellowish pigment. Rods : 0.5 to 0.7 by 1.25 to 2.5 microns. Motile with peritrichous flagella. Gram- negative. Gelatin colonies: After 3 days at 20°C, circular, 1 to 1.5 mm in diameter, yellow- ish-white, convex, entire. Microscopi- cally gray with opaque borders and darker patches. Gelatin stab : Surface growth somewhat umbonate. In 10 to 12 days a slow lique- faction. Intense yellow growth. Agar colonies: Grayish-white, fatty lustre, turning yellow after several days. Agar slant: Growth grayish-white, fatty lustre, becoming yellow. Broth : Strong more or less flocculent turbidity. No surface growth. Little sediment . Potato: Growth somewhat raised, be- coming yellowish. Milk: Coagulated in 14 days, becoming alkaline, slowly clearing. Indole is formed. Nitrites not produced from nitrates. Acid and gas from glucose, sucrose and mannitol. No gas from lactose and glycerol. Optimum temperature 20°C. Source : From cotton plants. Habitat : Causes a root-rot of cotton (Gossypium sp.). 12. Erwinia flavida (Fawcett) Magrou. {Bacillus flavidus Fawcett, Rev. Indust. y Agrico. de Tucuman, 13, 1922, 5; Rev. Xpp. Mycology, 2, 1923, 338; not Bacillus flavidus Morse, Jour. Inf. Dis., 11, 1912, 284; Magrou, in Hauduroy et al.. Diet, d. Bact. Path., 1937, 207.) From Latin flavus, yellow. Morphology : Motile with peritrichous flagella. Gram-negative. Gelatin: Yellow growth. Liquefac- tion. Milk: Coagulated. Potato: Yellow growth. Indole is formed. 472 MANUAL OF DETERMINATIVE BACTERIOLOGY Nitrites not produced from nitrates. Acid and gas from glucose, lactose and sucrose. Diastase not formed. Source : From sugar cane. Habitat : Causes a soft rot of sugar cane (Saccharum officinarum). Note : If this decay is due to a simple organism as stated above, it is probable that it should be considered merely a chromogenic strain of Erwinia carotovora. 13. Erwinia dissolvens (Rosen) comb, nov. {Pseudomonas dissolvens Rosen, Phytopath., 12, 1922, 497; Phytomonas dissolvens Rosen, Phytopath., 16, 1926, 264; Bacterium dissolvens Rosen, ibid.; Aplanobacter dissolvens Rosen, ibid.; Aerobacter dissolvens Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 473.) From Latin, dissolving. Rods: 0.5 to 0.9 by 0.7 to 1.2 microns. Pairs, rarely in chains. Capsules pres- ent. First described as motile with a single flagellum; later as non-motile. Gram-negative. Gelatin: Not liquefied. Agar colonies: Round, margins entire, white, opaque, glistening, butyrous, emitting a strong odor of decaying vege- tables. Broth : Turbid with heavy surface growth consisting of ring, and floccules or compact slimj^ masses and streamers. Abundant sediment. Uschinsky's solution: Good growth, but not viscid. Litmus milk: Acid, coagulated. Indole produced. Nitrites produced from nitrates. Hydrogen sulfide not produced. Acid and gas from glucose, galactose, mannitol, sucrose, maltose, lactose and glycerol. Starch hj'drolyzed. Optimum temperature 30°C. Good growth in 3 per cent salt. Re- tarded at 4 per cent. Source: From rotting corn stalks. Habitat : Pathogenic in corn plants. 14. Erwinia nimipressuralis Carter. (Illinois Nat. Hist. Survey Bull. 23, 1945, 423.) From Latin nimis, too much and pressuralis, pertaining to pressure. Rods: Mostly 0.34 to 0.68 by 0.68 to 1.35 microns. Motile with as many as 6 peritrichous flagella. Capsules not observed. Gram-negative. Gelatin: Not liquefied. Potato glucose agar : Colonies circular, smooth, whitish-cream, entire, flat to slightly raised and usually opaque. Gas produced when medium is stabbed. Broth : Abundant with thin pellicle or flocculent surface growth. Sediment scant and viscid. Gas produced in nutrient broth plus glucose was 47 per cent CO2 and 2.4 per cent hydrogen. CO2 varied with age of culture, more being produced in young cultures. Milk: Acid, coagulated. Litmus and bromocresol purple are reduced. Not peptonized. Nitrites produced from nitrates. Hydrogen sulfide produced. Indole not produced. Acid and gas produced from arabinose, rhamnose, xylose, glucose, fructose, galactose, mannose, lactose, maltose, trehalose, melibiose, cellobiose, manni- tol, sorbitol and salicin; no acid or gas from inulin, dextrin or filterpaper; variable results from sucrose, raffinose, melezitose, dulcitol, glycerol and elm sawdust. Pectin is not fermented. Starch not hydrolyzed. Methyl red test positive. Acetyl- methylcarbinol produced. Facultative anaerobe. Optimum temperature 24° to 30°C. Maximum 37°C. Minimum 5°C or lower. Thermal death point 45° to 55°C. Optimum pH for growth 6.8 to 7.5. Maximum 10.0+. Minimum 4.6. Source : Five cultures from 5 different trees affected with wet wood. Habitat : Pathogenic in trunk wood of elms, Ulmus americana, U. pumila, U. fulva and U. procera. FAMILY EXTEROBACTERIACEAE 473 15. Erwinia ananas Serrano. (Philip- pine Jour. Sci., 36, 1928, 271; Bacillus ananas Serrano, ibid.; Bacterium ananas Burgwitz, Phytopathogenic Bacteria, Leningrad, 1935, 44.) Named for the genus. Ananas. Note : Not to be confused with Pseudo- monas (Phytomonas) ananas Serrano, Philippine Jour. Sci., 36, 1928, 271. Short rods: 0.6 by 0.9 micron, with rounded ends, occurring singly, in pairs and in short chains. Encapsulated. Motile with peritrichous flagella. Gram- negative . Gelatin stab: Stratiform liquefaction, with a deep chrome-yellow sediment. Potato glucose agar: After 24 hours, circular, 3 mm in diameter, convex, dense, homogeneous, entire, moist, straw- yellow, mottled, becoming primuline yellow. Plates have a molasses odor. Show two types of colonies, rough and smooth. Rough colonies have crenate margins. Potato glucose agar slant : Growth straw-yellow, raised, becoming primu- line yellow, moist, glistening. Broth: Turbid, with a straw-colored pellicle and ring. Glucose broth: Growth sulfur yellow. Litmus milk: Coagulated, faintly acid, becoming alkaline. Potato : Copious growth, moist, glisten- ing, spreading, becoming primuline yel- low. Indole not formed. Blood serum : Moderate growth, slightly raised, mustard yellow to primu- line yellow. No liquefaction after 3 months. Cohn's solution: No growth. Phenol negative. Diastase produced. Nitrites produced from nitrates. Slight amount of ammonia produced. Slight amount of H2S produced. Small amount of alcohol and aldehyde produced. No gas from carbohydrates. Acid from glucose, lactose, sucrose, mannitol, raffinose, glycerol, salicin, dextrin, mal- tose, fructose and mannose. No acid from arabinose, xylose, amygdalin, rham- nose, inositol, inulin, dulcitol, adonitol, asparagine or starch. Source : From the pineapple {Ananas sativum) and sugar-cane {Saccharum offi,- cinarum). Habitat : Causes a brown rot of the fruitlets of pineapple. 16. Erwinia cytolytica Chester. (Phy- topath., 28, 1938, 431.) From Latin, cell dissolving. Rods: 0.6 to 0.7 by 2.5 to 3.5 microns. Singly or in pairs. Gram-negative. Mo- tile with peritrichous flagella. Gelatin: Slow liquefaction. Agar colonies: 2 to 3 mm in diameter, round, convex, moist, glistening, grayish- white, watery, translucent. Light brownish-yellow by transmitted light. Broth: Turbid. Milk: Coagulated in 5 to 7 days. Slightly acid. Not digested. Nitrites produced from nitrates. Indole not formed. Hydrogen sulfide not formed. Acetylmethylcarbinol : A slight reac- tion. Acid without gas from glucose, lactose, sucrose, raffinose, mannitol, salicin and isodulcitol. No acid from fructose, arab- inose, xylose, glycerol and inulin. Starch hydrolyzed. Pectin dissolved. Asparagine, peptone, and ammonia used as nitrogen sources in synthetic medium plus glucose. Potassium nitrate not used. Optimum temperature 28° to 30°C. Growth at 37°C. Slow growth at 20°C and no growth at 8° to 10°C. Good growth at pH 6.8 to 7.3. Feeble growth at 5.0. No growth at 4.4, Aerobic and facultative anaerobic. Source : Several isolates from diseased dahlias in New York Botanical Garden. Habitat : Causes a rot of the tuber and stems of dahlias. 474 MANUAL OF DETERMINATIVE BACTERIOLOGY 17. Erwinia aroideae (Towusend) Hol- land. {Bacillus aroideae Townsend, U. S. Dept. Agr., Bur. Plant Ind. Bull. 60, 1904, 40; Holland, Jour. Bact., 5, 1920, 222; Bacterium aroideae Stapp, in Sor- auer, Handb. d. Pflanzenkr., 5 Aufl., 2, 1928, 41 ; Pectobacterium aroideae Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 472.) From Greek, pertaining to the family Araceae. Probable synonyms : Erwinia croci (Mizusawa) IVIagrou. {Bacillus croci Mizusawa, Kanag. Agr. Exp. Sta. Bull. 51, 1921, 1; Ann. Phytopath. Soc. Japan, 1, 1923, 1; Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 204.) Attacks Crocus saiivus, also onion. Erwinia melonis (Giddings) Holland. {Bacillus melonis Giddings, Vermont Agr. E.xp. Sta. Bull. 148, 1910, 413; Hol- land, Jour. Bact., 5, 1920, 222; Pectobac- terium melonis Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 473.) E. F. Smith (An Introduction to Bact. Dis. of Plants, 1920, 240) considered Erwinia melonis and Erwinia aroideae identical. Causes a soft rot of muskmelon. Erwinia papaveris (Ayyar) Magrou. (Bacillus papaveris Ayyar, Mem. Dept. Agr. India, Bact. Ser. 2, 1927, 29 ; Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 214.) The cause of a soft rot of the garden poppy. Rods: 0.5 by 2 to 3 microns, with rounded ends, occurring singly, in pairs and in fours, also in chains under certain conditions. Motile with peritrichous flagella. No capsules. Gram-negative. Gelatin stab : Narrow infundibuliform liquefaction. Agar colonies: Circular to amoeboid, white, glistening. Borders sharp. Gran- ular structure. Agar slant : Growth white to grayish- white, moist, glistening. Medium not discolored. Broth: Turbid. Potato: Growth whitish, with tinge of yellow. Medium graj^ed. Litmus milk: Coagulated, acid, with separation of whey, not peptonized. Litmus reduced. Indole not formed. Nitrites produced from nitrates. Acetylmethylcarbinol produced. IMethyl red negative (Dowson, Cent. f. Bakt., II Abt., 100, 1939, 183). Acid without gas from glucose, lactose, sucrose, maltose, mannitol, glycerol, fructose, raffinose, arabinose and xylose. Growth in closed arm. Diastase slight. Hydrogen sulfide produced. Uschinsky's solution: Good growth. No growth in nitrogen. Growth feeble in Ho and CO2. Temperature relations : Optimum 35 °C. Minimum 6°C. Maximum 41 °C. Ther- mal death point 50°C for 10 minutes. Facultative anaerobe. Differential characters: See Erwinia carotovora. INIassey (Phytopath., 14, 1924, 460) considered Erwinia aroideae and Erwinia carotovora distinct species, though closely related. Link and Talia- ferro (Bot. Gazette, 85, 1928, 198) found them distinct serologically. Dowson (Ann. Appl. Biol., 28, 1941, 102) differen- tiated them on their action on maltose and xylose. Source : From rotted calla lily. Habitat : Causes a soft rot of calla. Affects raw potato, egg-plant, cauli- flower, radish, cucumber, cabbage, pars- nip, turnip, salsify, tomato (ripe and green). 18. Erwinia mangiferae (Doidge) Ber- gey et al. {Bacillus mangiferae Doidge, Ann. Appl. Biol., ^, 1915, 1; Bergeyetal., iManual, 1st ed., 1923, 173.) Named for the genus, Mangifera. P^ods: 0.6 by 1.5 microns, occurring singly and in chains, with rounded ends. Encapsulated. Motile with peritrichous flagella. Gram-negative. Gelatin stab : Medium liquefied in 10 to 17 days. Growth yellow. Agar colonies: Glistening, yellowish, undulate borders. FAMILY EXTEROBACTERIACEAE 475 Agar slant : Growth yellow, glistening. Broth : Turbid, with yellow ring. Litmus milk: Slow coagulation at 37 °C. Slight acidity. Casein slowly dissolved. Litmus reduced. Potato : Growth spreading, glistening, yellowish. Medium not discolored. Indole formed in peptone solution. Phenol negative. Nitrites produced from nitrates. No HoS produced. No ammonia in broth. Feeble acid production without gas from glucose, lactose, sucrose, fructose and glycerol. No growth in closed arm with lactose and glycerol; more or less growth in closed arm with glucose, su- crose, fructose, maltose, raffinose and mannitol. Diastase not formed. Produces an enzyme capable of dis- solving the middle lamella but without action on cellulose. Cohn's solution: Slight turbidity. Usehinsky's solution : No growth. Fermis' solution with starch jelly : No growth. Pigment insoluble in water, alcohol, ether, chloroform or dilute acids. Temperature relations : Optimum 30°C. IMinimum 5° to 6°C. Maximum 45°C. Thermal death point 60°C. Source : From the mango in Africa. Habitat : Causes a disease of the mango (Ma n gife ra indica) . 19. Erwinia citrimaculans (Doidge) Magrou. (Bacillus citrhnaculaiis Doidge, Ann. Appl. Biol., 3, 1917, 53; Bacterium citrimaculans Burgwitz, Phytopath. Bac- teria, Leningrad, 1935, 154; INIagrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 203.) From the genus Citrus and Latin 7uaculans, spotting. Rods: 0.45 to 0.7 by 0.8 to 3.2 microns. Motile with peritrichous flagella. Con- spicuous capsule present. Gram-posi- tive. Dowson thinks this species Gram- negative (Cent. f. Bakt., II Abt., 100, 1939, 184). Gelatin : Liquefied. Agar colonies: Subcircular, j-ellow, with dense grumose centers. Broth: Turbid, with pellicle and sedi- ment. ]Milk : Coagulated, with precipitation of casein and extrusion of whey. Not peptonized. Litmus gradually reduced. Blood serum : Not liquefied. Indole is formed. Nitrites produced from nitrates with evolution of gas. Ammonia produced in broth. Acid without visible gas from glucose, sucrose, fructose, galactose, maltose and mannitol. No acid from lactose, glyc- erol, dextrin or starch. Diastase not produced. Cohn's solution: No growth. Usehinsky's solution: Growth present. No growth in broth over chloroform. Methylene blue and neutral red re- duced. Pigment insoluble in water, alcohol, ether, chloroform, carbon bisulfide, dilute acid or alkalis. A turbid growth is produced in 10 per cent salt. Temperature relations : Optimum 35°C. ^Maximum 43°C. Thermal death point 62°C for 10 minutes. Facultative anaerobe. Source : From diseased lemons and oranges. Habitat : Causes a spot disease of citrus. In nature attacks lemons, or- anges, naartjes and has also been success- full}^ inoculated into limes, shaddock, grapefruit and citron. Seville oranges are resistant. 20. Erwinia rhapontici (^Millard) comb. 710V. (Phytomonas rhapontica Millard, Univ. Leeds and Yorkshire Council for Agr. Ed. Bui. 134, 1924, 111; Bacterium rhaponticum Millard, ibid.; Aplanobacter rhaponticum Elliott, Man. Bact. Plant Path., 1930, 12.) From Greek Rha pontic, rhubarb of Pontus, a province of Asia Minor; M. L. Rheum rhaponticum. 476 MANUAL OF DETERMINATIVE BACTERIOLOGY Description from Metcalfe, Ann. of Appl. Biol., ^7, 1940, 502, where he sug- gests it belongs in Erivinia. Rods: 0.5 to 0.8 by 1.2 to 1.5 microns. Motile with 3 to 7 peritrichous flagella. Gram-negative. Gelatin stab : Beaded growth. No liquefaction. Infusion agar: Colonies circular, con- vex, smooth, glistening, translucent, with margins entire, 2 to 3 mm in diam- eter in 48 hours at 25°C. Rhubarb agar : Colonies slightly larger, often with a yellowish tinge. Tryptophane broth : Turbid with frag- ile pellicle, a slight rim and slight floccu- lent deposit. Milk : Acid in 3 to 4 days with or with- out slight curd separation. No clotting. Indole not produced. Nitrites formed from nitrates. Acetylmethylcarbinol produced. No hydrogen sulfide produced. Cohn's solution : Moderate growth. Acid but no gas from arabinose, xylose, glucose, galactose, fructose, mannose, lactose, maltose, sucrose, mannitol, glyc- erol and salicin. Growth in citrate solution. Starch not hydrolyzed. Chromogenesis : Water-soluble pinkish pigment in various media. Growth from 0°C to 37 °C and possibly higher. Distinctive characters : Differs from Erwinia aroideae in that it does not liquify gelatin nor clot milk and is chro- mogenic . It also has a limited host range . Source : From rotting rhubarb crowns. Metcalfe used 6 isolates from various sources in describing the pathogen. Habitat : Causes a crown-rot of rhu- barb. 21. Erwinia lathyri (Manns and Tau- benhaus) Holland. {Bacillus lathyri Manns and Taubenhaus, Gardener's Chronicle, 53, 1913, 215; Manns, Dela- ware Agr. Exp. Sta., Bui. 108, 1915, 23; Holland, Jour. Bact., 5, 1920, 218; Bac- terium lathyri Burgwitz, Phytopath. Bacteria, Leningrad, 1935, 76.) From the genus LatMjrus. Rods : After 24 hours at 25° to 28°C, 0.6 to 0.85 by 0.75 to 1.5 microns, with rounded ends. No capsules. Motile with peritrichous flagella. Gram-nega- tive. Gelatin colonies: After 8 days, cir- cular, slightly convex, edges smooth. Liquefaction too slow to show on plate. Gelatin stab: Growth best at surface. Line of stab filiform. Liquefaction slow, fairly well begun in four weeks, complete in three months. Agar colonies: After 24 hours, yellow, stellate to amoeboid, smooth, glistening, slightly raised, entire. Centers gran- ular, yellow. Agar slant: Growth filiform, slightly convex, smooth, glistening, opaque, buty- rous, light to deep yellow. Odor absent. Broth: Strong turbidity in 24 hours, little or no pellicle. Sediment scant. Litmus milk: Slow increase of acidity, not always sufficient to cause coagulation. Digestion of casein slow and variable. Potato : Growth rapid, filiform, slightly convex, smooth, glistening, butyrous to slightly viscid. Light to deep yellow. Medium not discolored. Indole is formed. Cohn's solution: No growth. Uschinsky's solution: Rapid growth, sometimes a pellicle. Fluid viscid. Asparagine solution : Good growth. Nitrites are not produced from ni- trates. Ammonia produced in broth and as- paragine solution. No gas from carbohydrates. Acid from glucose, lactose, sucrose, mannitol and glycerol. No growth in closed arm. Diastase not formed or extremely weak. Growth in broth over chloroform absent. Growth inhibited by 4 per cent NaCl. Temperature relations : Optimum 28° FAMILY ENTEROBACTERIACEAE 477 to 30°C. Thermal death point 46° to 4S°C for 10 minutes. Aerobic. Source : From sweet peas. Habitat : Stated to be pathogenic for sweet pea {Lathyrus odoratus) and other legumes. Considered by many to be a saprophyte . 22. Erwinia lilii (Uyeda) Magrou. {Bacillus lilii Uyeda, by Bokura, Ann. Phytopath. Soc. Japan, 1{2), 1919, 36; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 210.) From Latin lilnim, a name taken from the Greek but derived from the Celtic word li meaning white; M. L., generic name, Lilium. Translated by ]\Iarion Okimoto. Rods: 0.6 to 0.7 by 0.8 to 1.0 micron. No capsules. Motile with 6 to 8 peri- trichous flagella. Gram-positive(?). Gelatin : Liquefaction. Gelatin plate : Colonies after 2 days, round and smooth with grayish surface. Broth: Alkaline, ammonia produced. Milk: Curd formation. Indole produced. Nitrites produced from nitrates. Hydrogen sulfide produced. Sugar medium changes to a brown color. Conjac not utilized. Aerobic, facultative. Optimum temperature 32° to 34°C. Killed in 3 min. at 50°C. Resists— 20°C for 30 min. Source : From brown spots on lily bulbs in Japan. Habitat : Causes a disease of lily bulbs and leaves. Appendix: The following additional species are found in the literature. Many are incompletely described. Bacillus hrassicaevorus Delacroix. (Compt. rend. Acad. Sci., Paris, HO, 1905, 1356.) Presumably causes a rot of cabbage. Bacillus farneiianus Pavarino. (Atti R. Accad. Naz. Lincei Rend. CI. Sci. Fis., Mat. e Nat., 20, 1911, 233.) Bacillus putrefaciens putridus Dela- croix. (Ann. Inst. Nat. Agron., 5, 1906, 154.) Pathogenic for tobacco. Bacillus solaniperda Migula. (Syst. d. Bakt., 2, 1900, 573; Bacillus krameri Chester, Man. Determ. Bact., 1901, 282.) Causes a soft rot of potato. Bacillus spieckermanni Jaczewski. (Elliott, Bacterial Plant Diseases, 1935, 67.) Name applied to a species de- scribed by Spieckermann (Landw. Jahrb., 31, 1902, 155) but left unnamed. Bacillus tabacivorus Delacroix. (Ann. Inst. Nat. Agron., 5, 1906, 266.) Said to cause collar rot of tobacco. Bacillus tabificans Delacroix. (Compt. rend. Acad. Sci., Paris, 137, 1903, 871.) Said to cause spotting of tobacco leaves. Bacterium loehnisi Kalantarian. (Kalantarian, Cent. f. Bakt., II Abt., 65, 1925, 301; Phytomonas loehnisii Bergey et al., Manual, 3rd ed., 1930, 276.) From diseased cotton plants. Peritrichous. Bacterium lycopersici Burgwitz . (Ztschr. f. Pflanzenkr., 34, 1924, 304.) From a blossom end rot of tomato. Erwinia alliariae (Omori) Magrou. {Bacillus alliariae Omori, Official Gaz. of Japan, 11, 1896, No. 3758; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 195.) Causes a root rot of horse- radish. Erwinia araliavora (Uyeda) Magrou. {Bacillus araliavorus Uyeda, Bull. Imp. Agr. Exp. Sta. Tokyo, 35, 1909, 61; Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 197.) Causes a root rot of ginseng. Erwinia asteracearum (Pavarino) Ma- grou. {Bacillus asteracearum Paravino, Atti R. Accad. Naz. Lincei Rend. CI. Sci. Fis., Mat. e Nat., Ser. 5, 21, 1912, 544; Magrou, in Hauduroy et al., Diet, d. Bact. Path., 1937, 199.) Pathogenic for the aster {Aster chinensis). Erwinia bussei (Migula) Magrou. (Ba- cillus B, Busse, Ztschr. f. Pflanzenkr., 7, 1897, 74; Bacillus bussei Migula, Syst. d. Bakt., 2, 1900, 779; Bacillus betae Leh- mann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 599; not Bacillus betae Migula, Syst. d. Bakt., 2, 1900, 779; Magrou, in 478 MANUAL OF DETERMINATIVE BACTERIOLOGY Hauduroy et al., Diet. d. Bact. Path., 1937, 200.) Pathogenic for the sugar beet. Erwinia cacticida (Johnston and Hitch- cock) Magrou. (B. cacticidus Johnston and Hitchcock, Trans, and Proc. Roy. Soc. South Australia, 47, 1923, 162; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 201.) Causes a rot of cactus. Envinia edgeworthiae (Hori and Bokura) Magrou. {Bacillus edge- icorthiac Hori and Bokura, Ideta Arata, Supplement to Handbook of the Plant Diseases of Japan, 1, 1925, 32; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 206.) Pathogenic on Edgeworthia chrysantha, an oriental shrub. Erwinia ixiae (Severini) Magrou. (Bacillus ixiae Severini, Annali di Botanica, Rome, 11, 1913, 413; Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 208.) Pathogenic on gladiolus and other bulbs. Erwinia nelliae (Welles) Magrou. (Ba- cillus nelliae Welles, Philippine Jour. Sci., 30, 1922, 279; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 213.) Erwinia papayae (Rant) Magrou. (Bacillus papayae Rant, Cent. f. Bakt., II Abt., 84, 1931, 483; Magrou, in Haudu- roy et al.. Diet. d. Bact. Path., 1937, 214.) Pathogenic on papaya. Erwinia sacchari Roldan. (Philippine Agrie., £0, 1931, 256; Bacillus saccharuin Roldan, idem; not Bacillus sacchari Janse, Mededeel. uit's Lands. Planten- tuin, 9, 1891, 1.) Erwinia scabiegcna (von Faber) Ma- grou. (Bacterium scabiegenum von Fa- ber, Arb. Kais. Biol. Anst. f. Land. u. Forstw., 5, 1907, 347 ; Bacillus scabiegenus Stapp, in Sorauer, Handb. d. Pfianzenkr., 5 Aufl., 2, 1928, 103 ; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 217.) Pathogenic for the beet (Beta vulgaris). Erwinia serbinowi (Potebnia) Magrou. (Bacterium beticola Serbinow, Zhurnal Bolezni Rastenii, 7, 1913, 237; not Bac- terimn beticola Smith, Brown and Town- send, Bur. Plant Ind., U. S. Dept. Agr., Bui. 213, 1911, 194; Bacterium serbinowi Potebnia, Kharkov Prov. Agr. Exp. Sta., 1, 1915, 1; Bacillus beticola Stapp, in Sorauer, Handb. d. Pfianzenkr., 5 Aufl., 2, 1928, 93; Bacillus serbinowi Elliott, Man. of Baet. Plant Pathogens, 1930, 75; Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 217.) Pathogenic for the sugar beet. Erwinia uvae (Kruse) Magrou. (Ba- cillo della bacterosidei grappoli della vite, Cugini and Macchiati, Le Stazione sperimentali ital., 20, 1891, 579; Bacillus uvae Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufl., 2, 1896, 329; Bacterium vvae Chester, Ann. Rept. Del. Agr. Exp. Sta., .9, 1897, 53 and 127; Magrou, in Hauduroy et al., Diet. d. Baet. Path., 1937, 220.) Pathogenic for the grape. Erwinia vitawra du Plessis syn. Clos- tridium baccarinii Bergey et al., Manual, 1st ed., 1923, 328. FAMILY ENTEROBACTERIACEAE 479 TRIBE III. SERRATEAE BERGEY, BREED AND MURRAY. (Preprint, Manual, 5th ed., October, 1938, vi.) Small, aerobic rods, usually producing a bright red or pink pigment on agar and gelatin. There is a single genus. Genus I . Serratia Bizio emend. Breed and Breed* (Bizio, Biblioteca italiana o sia Giornale de lettera, scienze e arti, 30, 1823, 288; Zaogalactina Sette, Sull'arrossimento straordinario di alcune sostanze alimentose osservato nella provincia di Padova I'anno 1819. Venezia, 1824, 51 ; Coccobacterium Schmidt and Weis, Die Bakterien, 1902, 10; Erythrobacillus Fortineau, Compt. rend. Soc. Biol., Paris, 58, 1905, 104; Dicrobactrum Enderlein, Sitzber. Gesell. Naturf. Freunde, Berlin, 1917, 309; Breed and Breed, Cent. f. Bakt., II Abt., 71, 1927, 435.) Named for Serafino Serrati, the Italian physicist who invented a steam boat at Florence before 1787. Small, aerobic, rapidly liquefying, nitrate reducing, Gram-negative, peritrichous rods which produce characteristic red pigments. White to rose-red strains that lack brilliant colors are common. Coagulate and digest milk. Liquefy blood serum. Typical species produce CO2 and frequently H2 from glucose and other sugars; also acetic, formic, succinic and lactic acids, acetylmethylcarbinol and 2,3 butylene glycol. Saprophytic on decaying plant or even animal materials. The type species is Serratia mnrcescens Bizio. Key to the species of genus Serratia. I. Pigment not especially water-soluble, readily soluble in alcohol. A. No visible gas from glucose. 1. Inconspicuous pellicle, if any, on plain gelatin. 1. Serratia marcescens. 2. Brilliant orange-red pellicle on plain gelatin. 2. Serratia indica. B. Produce enough H2 with the CO2 from glucose to show gas in fermentation tubes. 1. Acetylmethylcarbinol produced. 3. Serratia plymuthicum. 2. Acetylmethylcarbinol not produced. 4. Serratia kilensis. II. Pigment soluble in water and alcohol. 5. Serratia piscatorum. 1. Serratia marcescens Bizio. (Po- Sette, Memoria storico-naturale sulP lenta porporina, Biblioteca italiana, 30, arrossimento straordinario di alcune 1823,288.) From Latin, dissolving into a sostanze alimentose. Venezia, 8°, 1824, fluid or viscous matter. 51 ; Protococcus imetrophus Meneghini, Synonyms: Zoagalactina imetrofa 1838, see Trevisan, Rend. R. Inst. Lomb. * Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, Nov., 1937; further revision by Prof. Robert S. Breed, Nov., 1945. 480 MANUAL OF DETERMINATIVE BACTERIOLOGY di Sci. e Let., Ser. 2, 20, 1887, 797 ; Monas prodigiosa Ehrenberg, Bericht u. d. z. Bekanntmachung geeigneten Verhand- lungen d. Kgl. preuss. Acad. d. Wissen- schaften, 1849, 354; Palmella prodigiosa Montague, Bui. Soc. nat. et cent. d. agric. Paris, S^r. 2, 7, 1853, 527 ; Micraloa prodigiosa Zanardini, 1863, see Trevisan, loc. cit., 1887, 799; Bacteridium prodi- giosum Schroeter, in Cohn, Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 109; Micrococcus prodigiosus Cohn, ibid., 127; Bacillus prodigiosus Fliigge, Die Mikro- organismen, 1886, 284; Bacillus imetro- phus Trevisan, loc. cit., 797; Bacillus marcescens De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 976; Bacterium prodigiosurn Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 259; Liquidobacterium prodigiosurn Orla- Jensen, Cent. f. Bakt., II Abt., 22, 1909, 339 ; Erythrobacillus prodigiosus Winslow et al.. Jour. Bact., 5, 1920, 209; Dicro- bactrum prodigiosurn Enderlein, Bak- terien Cyclogenie, 1925, 279; Salmonella marcescens and Salmonella prodigiosa Pribram, Jour. Bact., 18, 1929, 384; Chromobacterium prodigiosurn Topley and Wilson, Principles of Bacteriology, 1, 1931, 402. Description largely taken from Breed and Breed, Jour. Bact., 9, 1924, 545. Short rods, sometimes almost spherical : 0.5 by 0.5 to 1.0 micron, occurring singly and occasionally in chains of 5 or 6 elements. Motile, with four peri- trichous flagella. Eight to ten flagella on cells grown at 20° to 25°C (De Rossi, Rivista d'Igiene, U, 1903, 000). Gram- negative. Gelatin colonies: Thin, slightly granu- lar, gray becoming red, circular, with slightly undulate margin. Liquefy the medium rather quickly. Gelatin stab: Infundibuliform lique- faction. Sediment in liquefied medium usually red on top, white in the depth. Agar colonies: Circular, thin, granular, white becoming red. R and S colonies with mucoid variants (Reed, Jour. Bact., 34, 1937, 255). Agar slant : White, smooth, moist layer, taking on an orange-red to fuchsin color in three or four days, sometimes with metallic luster. Broth : Turbid, may form a red ring at surface or slight pellicle, and gray sedi- ment. Litmus milk : Acid reaction with soft coagulum. A red surface growth de- velops. Little or no digestion takes place. Potato: At first a white line appears, which rapidly turns red. The growth is luxuriant and frequently shows a metallic luster. Produces acetic, formic, succinic and levolactic acid, ethyl alcohol, acetyl- methylcarbinol, 2,3 butylene glycol, CO2 and a trace of H2 from glucose (Pederson and Breed, Jour. Bact., 16, 1928, 183). Grows poorly or not at all in distilled water containing urea, potassium chloride and glucose. Indole not produced. Nitrites produced from nitrates. Formation of H2S : Produced from cys- teine, cystine or organic sulfur com- pounds containing either of these mole- cules. Produced from sulfur but not from sulfites, sulfates or thiosulfates (Tarr, Biochem. Jour., 27, 1933, 1869; 28, 1934, 192). Acetylmethylcarbinol is produced (Breed). Pigment soluble in alcohol, ether, chloroform, benzol and carbon bisulfide (Schneider, Arb. Bakt. Hochsch. Karls- ruhe, 1 , 1894, 210). Pigment may diffuse through the agar, i.e., shows solubility in water where strains are very deeply pigmented (Breed). Pigment not formed at 35°C. Sodium formate broth (Stark and England, Jour. Bact., 29, 1935, 26) : Cul- tures do not produce visible gas (Breed). Odor of trimethylamine is produced. Aerobic, facultative. FAMILY ENTEROBACTERIACEAE 481 Optimum temperature 25° to 30°C. No growth at 37°C. Source: Described by Bizio (Joe. cit.) and Sette {loc. cit.) from growth on corn meal mush (polenta). Habitat: Water, soil, milk, foods, silk worms and other insects. 2. Serratia indica (Eisenberg) Bergey et al. {Bacillus indicus Eisenberg, Bakt. Diag., 1 Aufl., 1886, 1; Bacillus indicus ruber Fliigge, Die Mikroorganis- men, 2 Aufi., 1886, 285; Micrococcus indicus Koch, Berichte ueber die Reise zur Erforschung der Cholera, 1887; Bacillus ruber indicus Kruse, in Fltigge, Die ]Mikroorganismen, 3 Aufi., 2, 1896, 302; Bacterium ruber indicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 112; Erylhrobacillus indicus Hol- land, Jour. Bact., 5, 1920, 218; Bergey et al.. Manual, 1st ed., 1923, 88; Breed and Breed, Jour, of Bact., 11, 1926, 76; Chromobacterium indicum Topley and Wilson, Princ. Bact. and Immun., 1, 1931, 402.) From Latin indicus, of India. Small rods: 0.5 by 1.0 to 1.5 microns. Motile with four peritrichous fiagella. Gram-negative. Gelatin colonies: Resemble those of Serratia marcescens. Gelatin stab : Liquefied rather quickl}'. Brilliant orange-red pellicle on plain gelatin. Agar colonies: Pink, with slightly serrate margin, spreading, with green irridescence. Agar slant : Luxuriant , dirty-white layer. Pigment produced best in alka- line media. Broth: Turbid, with white sediment. Litmus milk: Acid and coagulated. Digestion complete in 10 days. Potato : Luxuriant growth with or with- out pigment formation. Produces same products (except H2) from glucose as does Serratia marcescens (Pederson and Breed, Jour. Bact., 16, 1928, 183). Indole not produced. Nitrites produced from nitrates. Growiih with pigment production in distilled water containing urea, potassium chloride and glucose. Blood serum liquefied. Odor of trimethylamine. Sodium formate broth : Cultures do not produce visible gas (Breed). Pathogenic for laboratorj^ animals. Acetylmethylcarbinol is produced (Breed). Aerobic, facultative. Optimum temperature 25° to 35°C. No growth at 37°C. Cultures of this organism lose their ability to produce the orange-red pellicle on gelatin and then become practically indistinguishable from cultures of Ser- ratia marcescens. This would indicate that this so-called species is a rough strain of the former species (Breed). See Reed (Jour. Bact., S4, 1937, 255) for a discussion of dissociation phenomena in this genus. Source : Isolated from alimentary tract of a Java ape in India; also from milk can from Ithaca, N. Y. Habitat : Presumably widely dis- tributed. Apparently the following non-gelatin liquefying strain belongs with this spe- cies. Subcultures that are claimed to be derived from the original now liquefy gelatin. 2a. Serratia miquelii Bergey et al. (Named Bacillus ruber by Miquel and described in a letter to Hefferan, Cent, f . Bakt., II Abt., 11, 1903, 402; Erythro- bacillus ruber Holland, Jour. Bact., 5, 1920, 223; Bergey et al., Manual, 1st ed., 1923, 95.) Isolated from water by Miquel. 3. Serratia plymuthicum (Lehmann and Neumann) Bergey et al. (Roter 482 MANUAL OP DETERMINATIVE BACTERIOLOGY Bacillus aus Plymouth, Fischer, Zeitschr. f. Hyg., 2, 1887, 74; Bacterium ply- muthicum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 264; Bacillus plymouthensis Migula, Syst. d. Bakt., 2, 1900, 849; Erythrobacillus plymouthensis Holland, Jour. Bact., 5, 1920, 220; Bergey et al., IManual, 1st ed., 1923, 88.) Latin- ized from Plymouth, England. Distinct rods : 0.6 by 1 .5 to 2.0 microns with rounded ends, occurring singly and in short chains. Motile with peritrichous flagella. Gram-negative. Gelatin colonies : Like Serratia mar- cescens. Original culture mucoid. Gelatin stab : Crateriform liquefaction. Liquefaction as in Serratia niarcescens. Agar colonies : Like mucoid varieties of Serratia viarcescens . Agar slant : Sometimes show metallic luster. Pigment as in Serratia inarces- cens. Broth: Like Serratia marcescens. Litmus milk: Acid and coagulated. Potato: Growth violet pink, with or without metallic luster. Gas from glucose, lactose and sucrose, 70 to 80 per cent of it CO;. Remainder is Ho. Gas is also produced in asparagine solutions. Strong fecal odor produced. Blood serum liquefied. Acetylmethylcarbinol is produced (Breed). Sodium formate broth: Cultures pro- duce abundant gas (Breed). Pigment soluble in alcohol, ether and sometimes water. Aerobic, facultative. Optimum temperature 30 °C. Source : From water supply of PI3'- mouth, England. Habitat : Water and various foods. 4. Serratia kilensis (Lehmann and Neumann) Bergey et al. (Bacterium h, Breunig, Inaug. Diss., Kiel, 1888; Bacillus ruber hallicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 303; Bacterium kiliense Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 263 ; Bacterium, ruber balticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 142; Bacillus kiliensis Migula, Syst. d. Bakt., 2, 1900, 8i7 ; Enjthrobacil- lus kilieyisis Holland, Jour. Bact., 5, 1920, 218; Bergey et al.. Manual, 1st ed., 1923, 90 ; Chromobacterium kielense Topley and Wilson, Princip. Bact. and Immun., 1, 1931, 400.) From Kiel, a city on the Baltic Sea. Description taken from Kruse {loc. cit.) and Bergey et al. (loc. cit.). Slender rods: 0.7 to 0.8 by 2.5 to 5.0 microns, occurring singly. Motile with four peritrichous flagella. Gram-nega- tive. Deep gelatin colonies : Bright yellow. Gelatin liquefied slowly, usually becom- ing rose-red. Glucose gelatin stab: Rapid liquefac- tion. Occasional gas bubbles (Breed). Agar colonies: Small, red becoming magenta, smooth. Agar slant : Bright red becoming darker in old cultures. Agar stab : Turbid strongly pigmented water of condensation. Broth : Turbid. Usually reddened. Litmus milk : Acid ; at 20°C, coagulated slowly and pigment produced; at 35°C, coagulated rapidly and no pigment produced. Potato : Slight red growth, becoming luxuriant and darker. Indole not formed. Nitrites and free nitrogen produced from nitrates. Blood serum liquefied. Acid and gas from carbohydrates (Leh- mann and Neumann, loc. cit.). Gas from glucose, lactose and sucrose, 20 to 30 per cent of it CO2 (Bergey). Inactive lactic acid produced and not more than a trace of acetylmethylcarbinol or 2, 3 butylene glycol (Pederson and Breed, Jour. Bact., 16, 1928, 183). Sodium formate broth : Gas produced (Breed). FAMILY EXTEROBACTERIACEAE 483 Acetylmethylcarbinol not produced by the KrdI culture (Breed). Pigment formed at 37 °C. Pigment especially soluble in alcohol. Optimum temperature 30°C. -Aerobic. Distinctive characters: It is not cer- tain whether Breunig's original culture was a heavily pigmented strain of Serratia marcescens, or whether it was of the type described above. Cultures of both types have been widely distributed as the Kiel bacillus. Descriptions drawn up by Kruse (loc. cit.) and Lehmann and Neu- mann {loc. cit.) in 1896 state that this bacterium produces visible gas. while Migula in 1900 gives a description which fits Serratia marcescens. Moreover, cul- tures obtained under this name from various laboratories in Europe and Amer- ica are sometimes of one type and some- times of the other. As the Krdl culture distributed as Bacillus ruber balticus is widely known and has now been shown to differ from Serratia marcescens in that it is a distinct rod in ordinary media, forms visible gas from carbohydrates and even more abundant gas from sodium formate media, the name Serratia kilensis is used here for the Krdl culture. Serratia kilensis is a distinct rod like Serratia plymuthicum, but fails to produce acetyl- methylcarbinol. This use of the name Serratia kilensis given here also accords with the description drawn up by Bergey for the first edition of the Manual based on the study of a culture which he ob- tained many j-ears previously from Eu- rope (Breed). Source : From water at Kiel, Germany. Habitat : Presumably widely dis- tributed. 5. Serratia piscatonim (Lehmann and Neumann) Breed. (Microbe rouge de la sardine, Du Bois Saint-S^vrin, Ann. Inst. Past., 8, 1894, 155; Bacterium piscatorum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 263; Bacillus ruber sardi- nae Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufl., 2, 1896, 302 ; Bacterium ruber sardinae Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 112; Bacillus sardinae ^ligula, Syst. d. Bakt., 2, 1900, 852; Bacillus piscatorus Chester, Man. Determ. Bact., 1901, 257.) From Latin piscatorum, of fishermen. Short rods: 0.5 by 0.6 micron, occur- ring in pairs, sometimes in fours or (in broth) in long filaments. Actively mo- tile. Gram-negative. Gelatin colonies: Small, yellowish-gray becoming pink, very slimy. Carmine- red pellicle. Liquefaction. Gelatin stab : Rapid liquefaction. Grayish pellicle which becomes red after 24 hours and later precipitates. Slimy. Agar colonies: Dull, white to pinkish growth. Broth : Rapid turbidity. Thick, slimy, white pellicle which later turns red. Purplish sediment. Liquid becomes pink and syrupy. In old cultures the broth is brown. Potato: At 37° to 39°C, red pigment visible after 8 hours. At room tempera- tures growth is first white, slimy, later red. Strong odor of trimethylamine. Distinctive characters : Pigment sol- uble in alcohol, more soluble in water. Good pigment production at 37°C. Slimi- ness. Source : Isolated in 1893 from a box of oil-packed sardines at a canning-factory in France. Also found in the red pus from fishermen and sardine-factorj'- workers suffering from felons. In these lesions, this organism is associated with an anaerobe, but by itself it is not pathogenic. Habitat : Presumably widely dis- tributed. Appendix: Serratia marcescens has frequently been described under other names, particularly where brilliantly pigmented cultures have been found. Some of these and other related species are listed below. It is known that white 484 MANUAL OF DETERMINATIVE BACTERIOLOGY strains of these organisms occur in nature but these strains when found have prob- ably been placed in non-chromogenio genera of the family Enter ohacteriaceae. Bacillus ruber Frank. (Frank, in Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 3, 1875, 181; not Bacillus ruber Zimmer- mann, Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, 1, 1890, 24; not Bacillus ruber Miquel, see Cent. f. Bakt., II Abt., 11, 1903, 402; Bacterium ruber Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 113.) Grew in a warm place on rice cooked in chicken broth. Bacillus subkiliensis Petrow. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 273.) Dust contamination from air. Reported to resemble Bacillus kiliensis. Bacterium aurescens Parr. (Proc. Soc. Exp. Biol, and Med., 35, 1937, 563). A reddish-brown organism. This and the reddish-orange organism described by Tittsler (Jour. Bact., S3, 1937, 450), which are regarded as pigmented vari- ants of Escherichia coli, resemble the organisms in Serratia closely but do not liquefy gelatin. From water. Serratia amylorubra (Hefferan) Bergey et al. {Bacillus amyloruber Hefferan, Cent. f. Bakt., II Abt., 11, 1903, 313; Erythrobacillus amyloruber Holland, Jour. Bact., 5, 1920, 217 ; Bergey et al., Manual, 1st ed., 1923, 90.) From Mississippi River water and butternailk. Serratia esseyana Combe. (These, Ecole de M^d. Univ. Besangon, 1934, 1.) From well water at Essey . A study of an authentic culture shows this to be Ser- ratia marcescens (Breed). Serratia fuchsina (Boekhout and De Vries) Bergey et al. (Bacillus fuchsinus Boekhout and DeVries, Cent. f. Bakt., II Abt., 4, 1898, 497; Erythrobacillus fuchsinus Holland, Jour. Bact., 5, 1920, 218; Bergey et al.. Manual, 1st ed., 1923, 91.) Bacillus fuchsinus Migula. (Der rote Bacillus, Lustig, Diag. d. Bakterien d. Wassers, 1893, 72; Migula, Syst. d. Bakt., 2, 1900, 853.) Although these two organisms were named independently from different cultures, they were un- doubtedly identical. The original cul- tures of these species appear to have been heavy pigmented strains of Serratia marcescens showing a metallic luster. No authentic cultures are available. From water. Serratia gutturis Jan. (Bull. Soc. Sci. de Bretagne, 16, 1939, 34.) From spu- tum. Claimed to be different from Serratia marcescens on the ground that it will grow on an asparagine medium and that it reduces molybdates actively. Serratia marinorubra Zobell and Up- ham. (Bull. Scripps Inst. Oceanogra- phy, LaJolla, 5, 1944, 255.) From sea water. Grew only on sea water media when first isolated but later a culture studied by Breed (1944) became adapted to growth on ordinary media and then showed the characteristics of Serratia marcescens. Serratia miniacea (Zimmermann) Ber- gey et al. {Bacillus miniaceus Zimmer- mann, Die Bakterien unserer Trink- und Nutzwasser, Chemnitz, 1, 1890, 46; Erythrobacillus miniaceus Holland, Jour. Bact., 5, 1920, 219 ; Bergey et al.. Manual, 1st ed., 1923, 90.) Probably a heavily pigmented strain of Serratia marcescens or Serratia plymuthicum showing metallic luster. From water. Serratia pyoseptica (Fortineau) Ber- gey et al. {Erythrobacillus pyosepticus Fortineau, Thesis, Faculty of Medicine, Paris, 1904; abstract in Bull. Inst. Pas- teur, 3, 1905, 13; Bergey et al.. Manual, 1st ed., 1923, 89.) No constant differ- ences have been detected between Ser- ratia marcescens and authentic cultures of Serratia pyoseptica. From the shirt of a hospital patient. Pathogenic for guinea pigs and birds. Forms a soluble toxin. Serratia rubidaea Stapp. {Bacterium rubidaeum Stapp, Cent. f. Bakt., II Abt., 102, 1940, 251; ibid., 259.) From surface of plants and in composts. Char- acters much like those of Serratia marcescens. FAMILY ENTEROBACTERIACEAE 485 Serratia rutilescens (Hefferan) Bergey et al. (Bacillus rutilescens Hefferan, Cent. f. Bakt., II Abt., 11, 1903, 313; Erythrobacillus rutilescens Holland, Jour. Bact., 5, 1920, 220; Bergey et al.. Man- ual, 1st ed., 1923, 91.) The characters given do not distinguish this species from strains of Serratia marcescens that have nearly lost their power of pigment pro- duction except that it is reported to grow rapidly at 37°C. No authentic cultures appear to be available . From Mississippi River water. Serratia rutilis (Hefferan) Bergey et al. (Bacillus rutilis Hefferan, Cent. f. Bakt., II Abt., 11, 1903, 313; Erythro- bacillus rutilis Holland, Jour. Bact., 5, 1920, 220; Bergey et al., 1st ed., 1923, 94.) The original of this species appears to have been a heavily pigmented strain of Serratia marcescens or of Serratia ply- muthicum. No characters are given that distinguish it from these species and no cultures appear to be available. From Illinois River water. Serratia ster cor aria Jan. (Bull. Soc. Sci. de Bretagne, 16, 1939, 34.) From feces. Claimed to be different from Serratia marcescens because it attacks lactose, maltose and mannitol and reduces molybdates even more actively than Serratia gutturis. 486 MANUAL OF DETERMINATIVE BACTERIOLOGY TRIBE IV. PROTEAE CASTELLANI AND CHALMERS. (Manual of Trop. Med., 3rd ed., 1919, 932.) Ferments glucose but not lactose with formation of acid and usually visible gas. There is a single genus. Genus I. Proteus II a user.* (Hauser, Sitzber. d. phys.-med. Sozietat zu Erlangen, 1885, 156; Liquidobacterium Jensen, Cent. f. Bakt., II Abt., 32, 1909, 337; Spirilina Hueppe, Wiesbaden, 1886, 146; Eisenbergia Enderlein, Sitzber. Ges. Naturf. Freunde, Berlin, 1917, 315.) From Latin, having a changeable form. Straight rods. Gram-negative. Generally actively motile at 25°C, motility may be weak or absent at 37°C, peritrichous, occasionally very numerous flagella. Generally produce amoeboid colonies, swarming phenomenon, on moist medium. Marked pleo- morphism characteristic only of very young, actively swarming cultures. Ferment glucose and usually sucrose but not lactose. Three species in fermentable carbohy- drates produce small gas volumes even after prolonged incubation and an occasional culture does not produce gas. One species usually produces acid only. Urea de- composed and trimethylamine oxide reduced by all species. The type species is Proteus vulgaris Hauser. Key to the species of genus Proteus. I. No action on mannitol. A. Acid and gas from sucrose. 1. Acid and gas from maltose, a. Indole formed. 1. Proteus vulgaris. B. Acid and gas from sucrose (delayed). 1. No action on maltose. a. Indole not formed. 2. Proteus mirabilis. C. No action on sucrose (ordinarily). 1. No action on maltose. a. Indole formed. 3. Proteus morganii. II. Acid, occasionally a bubble of gas, from mannitol. A. Acid from sucrose (delayed). 1. No action on maltose. a. Indole formed. 4. Proteus rettgeri. 1. Proteus vulgaris Hauser. (Hauser, Trevisan, I generi e le specie delle Bat- Sitzungsber. d. phys.-mediz. Sozietat zu teriacee, 1889, 17; Bacterium vulgare Erlangen, 1885, 156; Bacillus proteus Lehmann and Neumann, Bakt. Diag., * Originally revised by Prof. M. W. Yale, New York State Experiment Station, Geneva, New York, Nov., 1938; revised by Prof. C. A. Stuart and Dr. Robert Rusti- gian, Brown University, Providence, Rhode Island, May, 1943. FAMILY ENTEROBACTERIACEAE 487 1 Aufl., 2, 1896, 243; Bacillus pruteus vulgaris Kruse, in Flligge, Die Mikro- organismen, 2, 1896, 272; Bacterium (Proteus) vulgaris Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 101; Bacillus vulgaris ]Migula, Syst. d. Bakt., 2, 1900, 707; Bacterium proteus anin- dologenes van Loghem, Ann. Inst. Past., 32, 1918, 295; Bacillus proteus-vulgaris Holland, Jour. Bact., 5, 1920, 220.) From Latin, common. Hauser described Proteus vulgaris as a rapid gelatin liquefier and Proteus mirabi- lis as a slow liquefier. VVenner and Rettger (Jour. Bact., 4, 1919, 332) found the property of liquefying gelatin too variable to serve as a basis for separation of species. They suggested that this differentiating character be set aside and the two species differentiated on the basis of maltose fermentation, the species fermenting the sugar receiving the name Proteus vulgaris and the species failing to attack it, Proteus mirabilis. This suggestion was accepted by Bergey et al.. Manual, 1st ed., 1923 and Weldin, Iowa Jour. Sci., 1, 1927, 147; and their work was confirmed by Rustigian and Stuart (Jour. Bact., 43, 1943, 198) and by Thorn- ton (Jour. Bact., 48, 1944, 123). Also see Moltke (Contributions to the Char- acterization and Systematic Classification of Bac. proteus vulgaris (Hauser), Levin and Munksgaard, Copenhagen, 1927, 156). Rods: 0.5 to 1.0 by 1.0 to 3.0 microns, occurring singly, in pairs and frequentlj- in long chains. Actively motile, with peritrichous flagella. Gram-negative. Gelatin colonies : Irregular, spreading., rapidly liquefying. Gelatin stab: Rapid, stratiform lique- faction. Agar colonies : Opaque, graj', spreading. Agar slant : Thin, bluish-gray, spread- ing over entire surface. Broth: Marked turbidity, usually with a thin pellicle. Litmus milk: Slightly acid, becoming markedly alkaline. Quick peptoniza- tion. Potato: Abundant, creamy to yellow- ish-gray growth, becoming brown. Indole formed. Nitrites produced from nitrates. Acetylmethylcarbinol not formed. Acid and gas from glucose, fructose, galactose, maltose and sucrose. Xo acid or gas from dextrin, lactose or mannitol. See Moltke (loc. cit.) for other fermenta- tion characters. Ratio Ho to CO2 is 1:1 (Speck and Stark, Jour. Bact., 44, 1942, 687). Putrefactive odor produced. Sodium citrate usually utilized as sole source of carbon. Formation of H2S : Produced from cysteine, cystine or organic sulfur com- pounds containing either of these mole- cules. Produced from sulfur and thio- sulfates (Tarr, Biochem. Jour., 27, 1933, 1869; 28, 1934, 192). Lead acetate turned brown. Aerobic, facultative. Optimum temperature 37 °C. Distinctive characters: X-Strains of Weil and Felix. Lehmann-X'eumann- Breed, Determinative Bact., Eng. Trans., 7th ed., 2, 1931, 493: "The discovery of proteus strains which may be aggluti- nated by tj'phus serum is of very great importance. These are the so-called X-strains from typhus patients found by Weil and Felix. Thej^ first cultivated strains X and X2 from the urine of typhus patients and later the famous X19. The two former were agglutinated weakly, the latter strongly (up to 1:50,000). The diagnosis of tj-phus bj' agglutination with strain X19 proved to be excellent and the reaction took place in the serum of almost 100 per cent of those suffering from the disease. . . . The typhus strains of pro- teus have recently been divided into the two types of Felix and Weil, the H forms and the O forms. The former grows as a thin opaque film, the latter lacks this character and grows as non-spreading slimy colonies; frequently without dis- 488 MANUAL OF DETERMINATIVE BACTERIOLOGY tinct flagella. . . ." (For further de- scription of H and O forms see Moltke, loc. cit.) The X2and X19 strains mostly ferment maltose. Source: From putrid meat, infusions and abscesses. Habitat : Putrefying materials. 2. Proteus mirabilis Hauser. (Hau- ser, Sitzungsber. d. phys.-mediz. Sozietat zu Erlangen, 1885, 156 ; Bacillus mirabilis Trevisan, I generi e le specie delle Bat- teriacee, 1889, 17; Bacillus proleus mira- bilis Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufl., 2, 1896, 276; Bacterium mirabilis Chester, Del. Coll. Agr. Exp. Sta., 9th Ann. Rept., 1897, 101; Bacillus pseudoramosus Migula, Syst. d. Bakt., 2, 1900, 817; not Bacillus pseudoramosus Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 441 ; Bacillus proieus-mirabilis Holland, Jour. Bact., 5, 1920, 220.) From Latin mirabilis, wonderful. Short rods: 0.5 to 0.6 by 1.0 to 3.0 microns, occurring singly, in pairs and frequently in long chains. Motile, pos- sessing peritrichous flagella. Gram-neg- ative. Gelatin colonies: Irregular, spreading. Gelatin stab : Slow, stratiform lique- faction. Agar colonies: Gray, irregular, spread- ing. Agar slant : Thin, bluish-gray, spread- ing over surface. Broth: Turbid, with thin gray pellicle and sediment. Litmus milk: Slightly acid, becoming alkaline, peptonized. Potato : Dirty -gray, spreading growth. Indole not formed. Acetylmethylcarbinol frequently pro- duced weakly. Nitrites produced from nitrates. Acid and gas from glucose, fructose and galactose. Acid and gas usually pro- duced slowly from sucrose. No acid or gas from lactose, maltose, dextrin or mannitol. The XK strains are mostly maltose negative. Putrefactive odor produced. Hydrogen sulfide is produced. Sodium citrate usually utilized as a sole source of carbon. Aerobic, facultative. Optimum temperature 37 °C. Source: From putrid meat, infusions and abscesses. Also reported as a cause of gastroenteritis (Cherry and Barnes, Amer. Jour. Pub. Health, 36, 1946, 484). Habitat : Putrefying materials. 3. Proteus morganii (Winslow et al.) Rauss. (Organism No. 1, Morgan, Brit. Med. Jour., 1, 1906, 908; Bacillus morgani Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 481; Bacterium morgani Holland, Jour. Bact., 5, 1920, 215; Bac- terium metacoli or Escherichia morgani Thj0tta, Jour. Inf. Dis., 43, 1928, 349; Salmonella morgani Castellani and Chal- mers, Man. Trop. Med., 1919, 939; Rauss, Jour. Path, and Bact., 42, 1936, 183; Morganella morganii Fulton, Jour. Bact., 46, 1943, 81 ; regarded by Fulton as the type species of the genus Morganella.) Named for Morgan, who first isolated this organism. Common name : Morgan's bacillus, type 1. Rods: 0.4 to 0.6 by 1.0 to 2.0 microns, occurring singly. Motile with peri- trichous flagella. See Rauss, loc. cit., for discussion of flagellation and relation to the swarming characteristic. Gram- negative. Gelatin colonies: Bluish-gray, homoge- neous, smooth, entire. Gelatin stab : No liquefaction. Agar colonies : Grayish or bluish-white, circular, entire. Agar slant : Grayish-white, smooth, glistening growth. Broth: Turbid. Litmus milk: Neutral, or becoming alkaline. Potato : Dirty-white, limited growth. FAMILY ENTEROBACTERIACEAE 489 Indole is formed. Nitrites are produced from nitrates. Acetylmethylcarbinol not formed. Acid and a small amount of gas from glucose, fructose, galactose and man- nose. Rarely from xylose. Does not attack lactose, sucrose, maltose, arabi- nose, raffinose, dextrin, salicin, mannitol, dulcitol, sorbitol, adonitol or inositol. Hydrogen sulfide not produced. Sodium citrate not utilized as sole source of carbon. Aerobic, facultative. Optimum temperature 37°C. Source: Isolated from the feces of infants with summer diarrhea. Habitat : In intestinal canal in normal or diarrheal stools. 4. Proteus rettgeri (Hadley et al.) Rustigian and Stuart. {Bacterium rett- geri Hadlej^ Elkins and Caldwell, Rhode Island Agr. Exp. Sta. Bull. 174, 1918, 169; Bacillus rettgeri St. John-Brooks and Rhodes, Jour. Path, and Bact., 26, 1923, 434; Eberthella rettgeri Bergey et al., Manual, 1st ed., 1923, 232; Shigella rettgeri Weldin, Iowa State College Jour. Sci., /, 1927, 181; Atypical enteric or- ganisms of the Shigella group. Cope and Kilander, Amer. Jour. Pub. Health, 32, 1942, 352; Proteus entericus Rustigian and Stuart, Jour. Bact., 45, 19i3, 198; Rustigian and Stuart, Proc. Soc. Exp. Biol, and Med., 53, 1943, 241.) Named for L. F. Rettger, the American bac- teriologist, who isolated this species in 1904. Rods: 0.5 to 0.8 micron long, occurring singly, in pairs and occasionally in chains. Usually non-motile at 37°C, but actively motile variants possessing peritrichous flagella can be obtained at 25 °C. Gram- negative. Gelatin colonies : Small, grayish, trans- lucent, entire. Gelatin stab: No liquefaction. Agar colonies: Small, grayish, translu- cent, entire; under suitable conditions some strains show marked spreading. Agar slant : Filiform to echinulate, grayish, thin, moist, translucent. Broth: Turbid with flocculent to viscid sediment. Litmus milk: Alkaline in eight days, becoming translucent. Potato: Luxuriant, grayish growth. Acid and occasionally slight gas from glucose, fructose, galactose and mannitol. Salicin may or may not be fermented. Slow and sometimes weak acid in sucrose. Lactose and maltose not fermented. Indole is formed. Nitrites are produced from nitrates. Acetj'lmethylcarbinol not formed. Hydrogen sulfide not produced. Sodium citrate utilized as sole source of carbon. Aerobic, facultative. Optimum temperature 37 °C. Source : Originallj' isolated from chol- era-like epidemic among chickens; re- cently isolated from sporadic and epi- demic gastroenteritis patients. Habitat : Fowl typhoid and some cholera-like diseases of birds. Appendix: Acceptance of gelatin lique- faction and fermentation of glucose and sucrose but not lactose as the cardinal characteristics of Proteus without ref- erence to urease production and small gas volumes has resulted in some cul- tures of Paracolobactrum (Borman et al.. Jour. Bact., 48, 1944, 361) being described as Proteus (Rustigian and Stuart, Jour. Bact., 49, 1945, 419). Included in the appendix are species of Proteus whose taxonomic position is not clear. Where descriptions permit, the probable taxo- nomic position of the organism is indi- cated. For purposes of reference, or- ganisms are also included which do not now merit species rank in the genus Proteus and organisms which will now be found in another genus. Bacillus agglomerans Beijerinck. (Botan. Zeitung, 4G, 1888, 740 or 749.) From nodules on the roots of red clover. Colonies like those of Proteus. 490 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacillus murisepticus pleomorphus Karlinski. (Karlinski, Cent. f. Bakt., 5, 1889, 193; Proteus of Karlinski, Stern- berg, Man. of Bact., 1893, 460.) From a urine discharge and from abscesses in the uterus. Sternberg regards this spe- cies as probably identical with Proteus vulgaris Hauser. Flarohacterium tneningilidis Hauduroy et al. (Bacillus luteus liqucfaciens Hau- duroy, Duhamel, Ehringer and Mondin, Compt. rend. Soc. Biol., Paris, 110, 1932, 362; Hauduroy et al., Diet. d. Bact. Path., 1937, 236.) Related to this species but differing in that it ferments lactose is the following: Bacterium coli var. luteoliquefaciens Lehmann and Levy, in Lehmann and Neumann, Bakt. Diag., 4 Aufi., 2, 1907, 34-t (Bacillus coli var. luteoliquefaciens Hauduroy, Du- hamel, Ehringer and Mondin, loc. cit., 1932, 363). Proteus alveicola Serbinow. (Jour. Microbiol., Petrograd, 2, 1915, 19.) From an infectious diarrhoea of hone.y bees (Apis mellifera). Proteus americanus Pacheco. (Scien- cia Medica, 6, 1928.) From the blood of patients with liver abscesses. Assis (Brasil Medico, No. 42-45, 1934, 35), St. John-Brooks and Rhodes (3rd Internat. Cong, for Microbiology, Rept. of Proc, 1939, 167), Rustigian and Stuart (Jour. Bact., 45, 1943, 198) and Thornton (Jour. Bact., 48, 1944, 123) agree that Proteus americanus is Proteus mirabilis. See Manual, 5th ed., 1939, 434 for a descrip- tion of this species. Proteus ammoniae Magath. (Magath, Jour. Inf. Dis., 43, 1928, 181 ; Salmonella ammoniae Hager and Magath, Jour. Amer. Med. Assn., 85, 1925, 1352.) From urine in cystitis. St. John-Brooks and Rhodes (3rd Internat. Congr. for Micro- biology, Rept. of Proc, 1939, 167), Levine (Jour. Bact., 43, 1942, 33), Rustigian and Stuart (Jour. Bact., 45, 1943, 198) and Thornton (Jour. Bact., 48, 1944, 123) agree that Proteus ammoniae is Proteus mirabilis. See Manual, 5th ed., 1939, 434 for a description of this species. See Fulton, Jour. Bact., 51, 1946, 685 for the view that Proteus ammoniae is a valid species. Proteus hombycis Bergey et al. (A Gram-negative bacillus, Glaser, Jour. Bact., 9, 1924, 344; Bacterium bombyci- vorum Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 445; Aerohacter hombycis Bergey et al.. Manual, 3rd ed., 1930, 334; Bergey et al., Manual, 4th ed., 1934, 365.) From diseased silk worms (Bombyx mori) . Proteus hombycis ap- pears to be a strain of Paracolobaclrum aerogenoides Borman et al. See Manual, 5th ed., 1939, 436 for a description of this species. Proteus diffluens (Castellani) Castel- lani and Chalmers. (Bacillus diffluens Castellani, 1915; Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 943.) From gastroenteritis patients. This may be a biochemical variant of Proteus mirabilis. Proteus henricensis Shaw. (Sci., 65, 1927, 477.) From putrefying materials. Said to be related to Proteus diffluens. Proteus infantum (Weldin and Levine) Weldin. (Dean, Med. Jour. Australia, 1, 1920, 27; Bacterium, infantum Weldin and Levine, Abst. Bact., 7, 1923, 13; Weldin, Iowa State Coll. Jour. Sci., 1, 1926, 148.) From urine and feces of an infant. Proteus inseclicolens Steinhaus. (Jour. Bact., 42, 1941, 763.) From the stom- ach of the milkweed bug (Oncopeltus fasciatus). This appears to be a strain of Paracolobactrum intermedium Borman et al. Proteus melanovogenes Miles and Hal- nan, (.lour. Hyg., 37, 1937, 79.) From eggs showing black rot. This does not appear to be a member of the genus Proteus. Proteus metadiffluens (Castellani) Cas- tellani and Chalmers. (Bacillus meta- diffluens Castellani, 1915; Castellani and Chalmers, Manual Trop. Med., 1919, 943.) From gastroenteritis patients. FAMILY EXTEKOBACTEKIACEAE 491 This does not appear to be a member of the genus Proteus. Proteus nadsonii Lobik. (Diseases of Plants, St. Petersburg, 9, 1915, 67.) From decomposed potatoes and toma- toes. This does not appear to be a mem- ber of the genus Proteus. Proteus noctuarum (White) Bergey at al. (Bacillus noctuarum White, Jour. Agr. Res., 26, 1923, 488; Escherichia noctuarii Bergey et al., IVIanual, 3rd ed., 1930, 327; Bergey et al., jVIanual, 4th ed., 1934, 363.) A cause of cutworm (Fam. Noctuidae) septicemia. Cultur- ally identical with but serologically different from Proteus sphingidis. Proteus odorans Pribram. {Bacterium aquatile odorans von Rigler, Hyg. Rund., 12, 1902, 479; Pribram, Klassifikation der Schizomyceten, Leipzig and Wien, 1933, 73.) From bottled mineral waters. Aromatic odor in milk. Proteus paraamericanus Magalhaes and Aragao. (Brasil Medico, 47, 1933, 815.) From urine. Assis (Brasil Medico, No. 42-45, 1934, 35) states that this is Proteus mirabilis. Proteus paradiffluens (Castellani) Cas- tellani and Chalmers. {Bacillus para- diffluens Castellani; Castellani and Chal- mers, Manual Trop. Med., 3rd ed., 1919, 943.) This appears to be identical with Proteus mirabilis. Proteus paramorganii Castellani and Chalmers. (Man. Trop. Med., 3rd ed., 1919, 943.) This is an H form of Proteus morganii. Proteus photuris Brown. (Amer. Mu- seum Nov., No. 251, 1927, 9.) From luminous organ of the firefly {Photuris pennsylvanicus) . This does not appear to be a member of the genus Proteus. Proteus piscicidus versicolor Babes and Riegler. (Babes and Riegler, Cent. f. Bakt., I Abt., Orig., 33, 1902-03, 449; Bacillus piscicidus versicolor Nepveux, These, Fac. Pharm., Paris, 1920, 114.) From diseased carp (Cyprinus carpio). Resembles Proteus vulgaris. Proteus pseudovaleriei Assis. (Jour. Hyg., 27, 1927, 108.) Rustigian and Stuart (Proc. Soc. Exper. Biol, and Med., 53, 1943, 241) state that this is a para- colon organism, presumably Paracolo- bactrum coliforme Borman et al. See Manual, 5th ed., 1939, 435 for a descrip- tion of this species. Proteus recticolens Steinhaus. (Jour. Bact., 42, 1941, 763.) From pylorus and rectum of the milkweed bug {Oncopeltus fasciatus). This appears to be a strain of Paracolobactrum intermedium Borman et al. Proteus sphingidis (White) Bergey et al. {Bacillus sphingidis White, Jour. Agr. Res., 26, 1923, 49; Escherichia sphingidis Bergey et al.. Manual, 3rd ed., 1930, 327; Bergey et al., Manual, 4th ed., 1934, 366.) A cause of hornworm septicemia {Protoparce sexta Johan. and P. quinquemaculata Haw.). See Manual, 5th ed., 1939, 605 for a descrip- tion of this species. White {loc. cit.) regards this species as possibly identical with Coccobacillus acridiorum d'Herelle. Proteus sulfureus Holschewnikoff. (Holschewnikoff, Fortschr. d. Med., 7, 1889, 201 and Ann. de Microgr., 1, 1888- 1889, 257; Bacillus lindenborni Trevisan, I generi e le specie delle Batteriacee, 1889, 17; Bacillus sidfureus Migula, Syst. d. Bakt., 2, 1900, 698; not Bacillus sulfureus Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From wa- ter. Similar to or perhaps identical with Proteus vulgaris Hauser. Produces H2S. Proteus sp. Steinhaus. (Jour. Bact., 42, 1941, 764.) This organism appears to be a strain of Paracolobactrum inter- medium Borman et al. Proteus sp. Warren and Lamb. (Jour. Med. Res., 44, 1924, 375.) From feces and blood of patient with a fatal infec- tion. This organism does not appear to be a member of the genus Proteus. Urobacillus liquefaciens septicus Krogius. (Compt. rend. Soc. Biol., Paris, 2, 1890, 65.) Regarded by Leh- mann and Neumann (Bakt. Diag., 1 Aufi., 2, 1896, 243) as a synonym of Pro- teus vulgaris. 492 MANUAL OF DETERMINATIVE BACTERIOLOGY TRIBE V. SALMONELLEAE BERGEY, BREED AND MURRAY. (Preprint, Manual, 5th ed., October, 1938, vi.) Rods that are either motile with peritrichous flagella or non-motile. Attack numerous carbohydrates with the formation of acid, or acid and gas. Lactose, su- crose and salicin are not ordinarily attacked. Do not produce acetylmethylcarbinol. Gelatin not liquefied (exceptions have been noted, but are rare). Urea not hydro- lyzed. Milk not peptonized. No spreading growth on ordinary 2 to 3 per cent agar. Live in the bodies of warm-blooded animals, including man, occasionally in reptiles, and frequently in food eaten by these animals. Key to the genera of tribe Salmonelleae. I. Ferments glucose with the formation of acid and, with few exceptions, gas. Genus I. Salmonella, p. 492. II. Ferments glucose with the formation of acid but. with rare exceptions, no gas. Genus II. Shigella, p. 535. Genus I. Salmonella Lignieres* (Rec. de m^d. v^t., S6r. 8, 7, 1900, 389.) Usuallj' motile, but non-motile forms occur. Produce acid and gas from glucose, maltose, mannitol and sorbitol (except that in Salmonella typhosa and S. gallinarum no gas is produced). Lactose, sucrose and salicin not attacked. Do not clot milk, form indole or liquefy gelatin. Reduce trimethylamine oxide to trimethylamine.f All of the known species are pathogenic for warm-blooded animals, including man, causing food infections and enteric fevers. A few are found in reptiles. Some or all may also live in decomposing foods. Although fermentation of lactose, sucrose and salicin, formation of indole, gelatin liquefaction and failure to produce gas have been described for organisms serologi- cally belonging to Salmonella, the practical recognition of this genus and studies of its constituent species suggest that these be looked upon as exceptions which do not invalidate the biochemical definition of the genus. Serological definition of the limits of the genus is fraught with many practical and theoretical difficulties. In- deed, there is increasing evidence of antigenic affinities of varying degree between Escherichia, Salmonella and Shigella. This is well reviewed by Bornstein (Jour. Immunol., 46, 1943, 439) . Within the limits of the genus Salmonella, serological rela- * Completely revised by Prof. Frederick Smith, McGill Univ., Montreal, P. Q., Canada, December, 1938; further revision, 1946. Manuscript read by Dr. F. KaufT- mann, State Serum Institute, Copenhagen, Denmark and by Dr. Philip Edwards and Dr. D. W. Bruner, Agri. Exper. Sta., Lexington, Kentucky, May, 1946. These spe- cialists have also assisted in completing references and in compiling records of the distribution of types. t Wood and Baird, Jour. Fish. Res. Board Canada, 6, 1943, 194. FAMILY ENTEROBACTERIACEAE 493 tionships are the chief means of identifying new strains. There is general dissatis- faction with the granting of species rank to each one of the rapidly mounting number of tj'pes. The purposes of the greater number of bacteriologists, however, will be best served for the present by listing the known types. There is a wide difference between the viewpoint of those who think of the sero- logical types recognized in this genus as species, e.g., Schiitze et al. (Jour. Hyg., 34, 1934, 333) and the more recently expressed viewpoint of Borman, Stuart and Wheeler (Jour. Bact., 48, 1944, 351). The latter authors recognize only three species in the genus. Salmonella choleraesuis, S. typhosa and S. kauffmannii. In the second report by Schutze et al. (Proc. 3rd Internat. Cong. Microbiol., New York, 1940, 832) the so- called species listed in the first report by this Sub-committee are designated as types. Kauffmann, who recognizes nearly 150 serotypes in the group, nevertheless notes in a recent paper (Acta Path, et Microbiol. Scand., 22, 1945, 144) that five types are of special interest in the field of human medicine. Salmonella paratyphi A, Salmonella paratyphi B, Salmonella paratyphi C, Salmonella typhi and Salmonella sendai; and that six types are of special interest in the field of veterinary medicine, Salmonella typhimurium, Salmonella abortusequi, Salmonella abortusovis, Salmonella cholerae- suis, Salmonella enteritidis and Salmonella gallinarum-pulloriim. The 150 or more serotypes are, in a way, comparable to the 50 or more serotypes of Diplococcus pneumoniae that are recognized on the basis of agglutination with im- mune serums. The serological methods used have proved to be of fundamental value as they provide useful diagnostic procedures by means of which unknown cultures can be accurately and quickly identified. As the morphology, staining properties and physiology of the bacteria belonging to the various types are practically identical, only the antigenic structure, source and habitat (so far as the latter is known) have been recorded for the majority of the types listed. Even though there is much duplication, descriptions similar to those used elsewhere in the Manual are given for the eleven types that are of greatest interest. Special mention has also been made of unusual characters such as failure to produce gas from glucose, lactose fermentation, indole production and gelatin liquefaction. The nomenclature used for this group presents a special problem. It developed from labelings used for cultures. These were designated by the name of a patient, e.g., Thompson; by the name of the hospital where the patient was placed, e.g., Vir- chow, Bispebjerg; or more frequently by the name of the village, locality or city where the outbreak occurred or was studied, e.g., Borbeck, Altendorf , Tel Aviv. The names of states and larger areas have also been used, e.g., Kentucky, Italia, etc. Recently several types have been named in honor of well-known bacteriologists, e.g., Berta, Gaminara, Arechavaleta. As this useful laboratory labeling is not in the form ordinarily used bj' taxonomists, various suggestions have been made regarding the development of a binomial nomenclature comparable to that more generally used. None of these suggestions has been generally accepted as yet. For example, Haupt (Ergebnisse d. Hyg., 13, 1932, 673) and others who have thought of the serotypes as species have added Latin endings to the place and other proper names that have been used, e.g.. Salmonella readingensis , S. rostockensis. Schutze et al. (Jour. Hj-g., 34, 1934, 333) accepted the view that the place and other names should be used in bi- nomials without adding Latin endings. Kauffmann (Ztschr. f . Hyg., 120, 1938, 193), on the other hand, has suggested that letters and numbers, e.g.. Salmonella B2, or even (Acta Path, et Microbiol. Scand., 22, 1945, 147) the antigenic formula be used with the generic name, e.g.. Salmonella IV, V, XII . . . b <-^ 1, 2 . . . instead o( Sal- monella paratyphi B. 494 MANUAL OF DETERMINATIVE BACTERIOLOGY The nomenclature used in the present edition of the Manual is slightly modified from that used in the fifth edition. The form adopted is in accordance with the view that the recognition of similar antigenic structures really identifies serotypes rather than species. In a way, serotypes are varieties in a taxonomic sense, though like horticultural varieties in higher plants, they do not exactly correspond with varieties as usually defined by taxonomists. Where cultural differences rather than antigenic structure have been used to subdivide species, these subdivisions are designated as varieties. As it is not clear as yet how many and what species will eventually be recognized, the form Salmonella sp. has been used as before to indicate that the serotypes belong to species in the genus Salmonella which are not yet definitely defined. Geographic and other proper names are used to designate types as these have been used exten- sively in the literature. They have an historic significance and are not as easily confused as are letters and numbers. No Latin endings have been used for these place names as this might indicate that the serotype names are accepted as species names. The genus Eberthella has been combined with the genus Salmonella as recommended by Schiitze et al. Hoc. cit.). With the exception of the typhoid organism, other species previously listed in Eberthella appear not to exist in type culture collections. As cultures are not available for study, these species are merely listed in an appendix to the genus Salmonella. The type species is Salmonella choleraesuis (Smith) Weldin. The table on pages 495 to 500 is used in place of the usual key. FAIVnLY EXTERCBACTERIACEAE 495 ^ re C«; X N C f,j s c^ c^i c-1 c^i :: c t c, ~ - cTcT"^ cTcrcririTs—'iiobZbC ^ eS l eS o « ^ x; y, y, y^y, y, y, yr^, y, y. y, y. y. y, y, >', y, y. y, y. y.y. y, X X > > > > > > > > > > > > > > > > > > > > > > > > > >.-^ i- ^ CO ~ O CC DQ CC aj ^ c X i^ o ^.- 5^^ >i%V — O 5 a, "^ « tc e p §,pi S,ti;QK250?Q=5-S< o c .^- :; '"Cx — ~xx — i; u 51. 5s E-i H H H H E-1 ?5« e ?a. B >>>>>>>>>>>>>>>>>>>>>> 1— 1 1— ( > > >>>>>>>>>>>>>>>>>>>> 1 la abortushovis la sp. (Type Bredeney) la sp. (Type Schleissheim) la sp. (Type Schwarzengrund) CJ «J CO ~ S;!5,j5,a,a,a,?i,si,=^f5,fi.?i,si,=i.=i,»,?i.?i.fi,a, •♦^COCOCOCOCOCOCCCOCQCCCOCOCOCOCOOOOOGQCO Sahnonel Salmonel Sahnonel Salmonel 0000000002222222222222 'c'c'B'e'e'o'a'e'c'c'c'e'e'e'a's'S'3'c'a'c's OQOQOSO^OQOQO^OQOQOQC^OQO^OQt^OQOQOSOQCQOoOQ ^ cq c<> CM ^ CI o (M CO i-HC<)CO-<+iiOOt^OOC10'-i(MeO-*iOc^t-OOC30 < ) ) m J FAMII.Y EXTKROBACTERIACEAE -497 >>>>>>>>>>>>> >>>>>>>>>f>>>> > > X X >< x" >< >< >< >< x" X X X x 2 s < C ~ M ,-> XI -- o ;:;'"■••• — o _^ M — a s c o .3 5.S h:; ^ i^ pq 3 - = -^ 5 -c 5b w ;2; H o < > ^ ;:q i: ^ ::; cK S c ;: H iHH-^HHHEHHH' ^ a, s, s. e s s B s 7-1 H H =0 00 si e e B 2 ^ c 9^ X x x o CJ o o o i >; >: >,; >> >; >" >! >, I ' ^. >. ^ H H H H H H H H H H H ^ S s ?i- ?5, SI, eeessBBs B c s s 4i ^ -. ^ ^ O 2 O O O CQ CC OQ CO <» '^ ^ o o o o Sees u u t^ ^ u u %> o s s s s s s o 498 MANUAL OF DETERMINATIVE BACTERIOLOGY c Si c < g bo E 0, CO CO t-ii-Hoo^i i>-(.-icrcrr-^ ,_( ,-4 _| ,_!,_, ^H 1—1 — T T— 1 I— 1 .— 1 t-H N s ■^ > '$■ > ii o,^'^ ^ _ — T — T -4" o q" oT o" aT tS -^ >> n sh C .5? 1 O _o o . in PiPP^>^PiP^p, ^Px><>> y. yr^r,^. y^ >i X X y. X X X X xxxxxxxxxxxxx f:4Cdbi^^f^b;i::^';df:^C4fd xxxxxxxxxxxxx Salmonella sp. (Type Eastbourne) Salmonella sp. (Type Panama) Salmonella sp. (Type Dar-es-Salaam) Salmonella sp. (Type Goettingen) Salmonella sp. (Type Java) Salmonella (jallinarum Salmonella pullorum Salmonella sp. (Type Canastel) Salmonella sp. (Type Italia) Salmonella sp. (Type Napoli) Salmonella sp. (Type Loma Jjinda) Salmonella sp. (Type New York) Salmonella sp. (Type London) Salmonella sp. (Type Give) Salmonella sp. (Type Uganda) Salmonella analis Salmonella sp. (Type Muenster) Salmonella sp. (Tyi)c Nyborg) Salmonella sp. (Tj'pe Vejle) Salmonella sp. (Type Meleagris) Salmonella sp. (T^q^e Shangani) Salmonella sp. (Type Zanzibar) Salmonella sp. (Type Amager) Salmonella sp. (Type Lexington) Salmonella sp. (Type Weltevreden) J5 t^OOOOOOQOajQOCOQOGOQOOl rH(Nec-*»ccot^oooo^(Meo OiOiOSOiOiOiCiOGtiOOOO 0 3 o c a w FAMILY ENTEROBACTERIACEAE 499 »0 lO CO t^ t^ lO 1 o 1 o 1 N 1 N cm" C C <0 "3 i-'i NT-,«r-l©i-H.-l^.-l.-l,-lr-(| I Sllrt ~. ~. ^ 2 CO > ^ ^ ^. III, X, XXVI III, X, XXVI III, XV III, XV III, XV (III), (XV), XXXIV I, III, XIX I, III, XIX I, III, XIX I, III, XIX (VIII), XX XI XI XI XI XI XI XI XIII, XXII, XIII, XXII XIII, XXII I, XIII, XXIII I, XIII, XXIIl I, XIII, XXIIl I, XIII, XXIII I, XIII, XXIII VI, XIV, XXIV Type Orion) Type Butantan) Type Newington) Tj^pe Selandia) Type New Brunswick) Type Illinois) Type Sonftenberg) Type Niloeso) Tjqie Sinisbury) Type Taksony) Typo Kentucky) Type Aberdeen) Type Rubislaw) Typo Pretoria) Tyi)o Venezia) Typo Solt) Type St. Lucie) Type Senegal) Type Marseille) Tyi)o Crunipy) Typ(> Poona) Typo Borbo(;k) Typo Mississippi) Typo Wichita) Ty])o Havana) Tyj)e Worthington) Typo Cuba) Type Ilcves) ji. Si, a ?i. Si, =^ CO CC CO CO cc sc e e e e e e s, a. ?i. ?s, CO eo CO CO e B e e oocoeoeocooooooocoeooococooocoeoooco monel monel monel monel monel monel monel monel monel monel monel monel monel monel monel monel monel hnonel huonel monel monel monel 'monel monel hnonel monel hnonel 'monel e c c « e c C<2 OQ Gq &5 &2 *^ c a e c CQ Cd cc CO ■* lO CO 1-- OC o o o o o o o O --I , 9i Ti !>iN^-D"C-^-C N,^ >3>%N SJ^e^-Tr^p^ a >■>■>■>■ to ^ ^X ><1 ^ c >y.y.y.y. < >< > > > > >_^ O ><^ ^ ►-H ^ HH >• • -H • — ^~ >^ ^ ;^ ^ is> > ►-> j:^ E > . . . . HH 1— 1 hH 1— ( 1— 1 ><^ > > > > P H S ^" . r J^ en > > > >■ >■ ^ y, y, y, y, y, y, y, y. y, ► . I-H l-H 1— t HH ^xxx'^;^;^^.^^x>^><:^'^>< o a , ^ - — - > >i >. >1 >i >,>> >V >!>.>> >i >. >. >i >, >•. >i >i ^>i bciSSoSbooobooboocooo pc« FAMILY ENTEKOBACTERIACEAE 5U1 1. Salmonella paratyphi (Kajser) Cas- tellani and Chalmers. {Bacterium -para- typhi Typus A, Brion and Kaj^ser, Munch, med. Wchnschr., 49, 1902, 611; Bacterium paratyphi Kayser, Cent. f. Bakt., I Abt., Orig., 31, 1902, 426; Ba- cillus paratyphosus A Boycott, Jour. Hyg., 6, 1906, 33; Bacillus paratyphi Winslow and Kligler, Jour. Bact., 1, 1916, 81; Bacillus paratyphosus Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 474; Salmonella paratyphi and Salmonella paratyphi A Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 938 and 939; Bacterium paraty- phosum A Holland, Jour. Bact., 5, 1920, 219.) From Latin para, like and typhus. typhoid. Rods: 0.6 by 3.0 to 4.0 microns, occur- ring singly. Motile with peritriohous flagella. Gram -negative. Gelatin colonies : Bluish-gray, homoge- neous, smooth, glistening, entire to slightly undulate. Gelatin stab : Fair surface grf)wth . Xo liquefaction. Agar colonies: Grayish, homogeneous, smooth, glistening, entire to slightly undulate. Agar slant: Filiform, grayish, smooth, glistening growth. Broth: Turbid, with slight grayish sediment. Litmus milk: Slightly acid. Potato: Limited, dirty-white streak. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, arabinose, maltose, trehalose, dextrin, glycerol, mannitol, dulcitol, rhamnose and sorbitol. No acid or gas from lactose, sucrose, raffi- nose, xylose, salicin, inulin, adonitol or inositol. Reduces trimethj-lamine oxide (Wood and Baird, Jour. Fish. Res. Bd. Canada, 6, 1943, 198). No hydrogen sulfide formed. Aerobic, facultative. Optimum temperature 37 °C. Antigenic structure: [I], II, XII: a: — . (Tj-pe Durazzo lacks I). Source: Isolated from enteric fever in man. Not known to be a natural patho- gen of animals. Habitat: A natural pathogen of man causing enteric fever. 2. Salmonella schottmuelleri (Winslow et al.) Bergej- et al. (Bacilli paraty- phique, Achard and Bensaude, Soc. M(^d. des Hop. de Paris, 13, 1896, 679; Bacillus paratyphi alcaligenes Schott- miiller, Deutsche med. Wchnschr., 32, 1900, 511; Bacterium paratyphi Typus B, Brion and Kayser, Miinch. med. Wchnschr., 49, 1902, 611; Bacillus para- typhosus B Boycott, Jour. Hyg., 6, 1906, 33; Bacterium paraiyphosum B Le Blaye and Guggenheim, Manuel Pratique de Diag. Bact., 1914; Bacillus schottmulleri Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 479; Salmonella paratyphi B Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Bacterium schottmiillcri Holland, Jour. Bact., 5, 1920, 222; included in Group IV of Hecht- Johansen, Copenhagen, 1923; Bergey et al., Manual, 1st ed., 1923, 213.) Named for Prof. Schottmuller who isolated this organism in 1899. Rods: 0.6 to 0.7 by 2.0 to 3.0 microns, occurring singly and in pairs. Motile with peritrichous flagella. Gram-nega- tive. Gelatin stab: No liquefaction. Agar colonies: Small, circular, bluish- gray, transparent, homogeneous, entire to undulate. Broth: Turbid with thin gray pellicle and sediment. Fecal odor. Litmus milk: Slightly acid, becoming alkaline. Potato: Grayish-white, viscous growth. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, arabinose, xylose, maltose, dextrin, trehalose, glycerol, 502 MANUAL OF DETERMINATIVE BAPTERIOLOGY mamiitol, dulcitol, sorbitol, rhamnose and inositol. No acid or gas from lac- tose, sucrose, inulin, salicin or adonitol and usually not from raffinose. Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide produced. Optimum temperature 37 °C. Aerobic, facultative. Antigenic structure: [I], IV, [V],XII: b: [1,2]. . . . Some strains lack antigen V and some have I. Source: Isolated from cases of enteric fever in man. Not a natural pathogen of animals. Habitat: A natural pathogen of man causing enteric fever. Also found rarely in cattle, sheep, swine, lower primates and chickens. 3. Salmonella sp. (Type Abony). {Sal- monella abony Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940, 1.) Antigenic structure: [I], IV, V, XII: b: e, n, X. . . . Source: Isolated by Kauffmann from a mixed culture of Salmonella abortus bovis sent to him by Dr. K. Rauss, Buda- pest. Later three additional cultures were received from Dr. Rauss. Original culture from the feces of a normal person . Habitat: All cultures thus far recog- nized have been from human sources. 4. Salmonella typhimurium (LoefRer) Castellani and Chalmers. {Bacillus ty- phi viurium Loeffler, Cent. f. Bakt., 11, 1892, 192; Bacterium typhi murium Ches- ter, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 70; Bacillus murium Migula, Syst. d. Bakt., 2, 1900, 761; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Bacillus typhi-murium Holland, Jour. Bact., 5, 1920, 221; Bac- terium typhi-murium Holland, idem; Bacillus enteritidis B, Typ. murium Januschke, Ztschr. f. Infektionskr. d. Haustiere, 27, 1924, 182.) The following are regarded as syn- onyms of this organism: Salmonella psittacosis Castellani and Chalmers (Man. Trop. Med., 3rd ed., 1919, 939; Bacillus psittacosis Nocard, Conseil d. Ilyg. Publique et Salubrity du Dept. du Seine, Stance, March 24, 1893; Bac- terium psittacosis Le Blaye and Guggen- heim, Manuel Pratique de Diagnostic Bacteriologique, 1914); Salmonella aer- trycke Castellani and Chalmers (Man. Trop. Med., 3rd ed., 1919, 939; Bacillus aertrycke De Nobele, Ann. Soc. Med. Gand., 72, 1898, 281; Bacillus para- aertrycke Castellani, Ann. di Med. Nav. e Colon., 11, 1914, 453; Bacterium aer- trycke Weldin and Levine, Abst. Bact., 7, 1923, 13) ; Kaensche's Bacillus and Basenau's Bacillus, Kaensche, Ztschr. f. Ilyg., 22, 1896, 53; Bacillus pestis- caviae Wherry (Jour. Inf. Dis., 5, 1908, 519; Bacillus cholera-caviae Wherry, Pub. Health Repts., November, 1908; Pasturella pestis-caviae Holland, Jour. Bact., 5, 1920, 219); Bacillus paratypho- sus B, Mutton type, Schtitze, Lancet, 1, 1920, 93; Group VII of Hecht-Johansen, Coi^enhagen, 1923; Salmonella aertrycke Ibrahim and Schiitze, Brit. Jour. Exp. Path., 9, 1928, 353; Bacterium enteritidis Breslau and Salmonella breslau of Ger- man literature; IMouse-typhoid of many authors. Some strains are confused with Salmonella anatis because of their origin in ducklings, e.g., see Salmonella ana- tum var. aertrycke Olsen and Goetchins, Cornell Vet., 27, 1937, 354. Hauduroy et al. (Diet. d. Bact. Path., Paris, 1937, 449) regard the following as synonyms of Salmonella aertrycke: Ba- cillus breslaviensis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 377; Bacterium breslaviensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 69; B. enteritidis breslaviense Berge, Deut. tierarztl. Wchnschr., 1926, 473; Salmonella meleagridis Rettger, Plas- tridge and Cameron, Jour. Inf. Dis., 53, 1933, 279; Salmonella aertrycke var. meleagridis Cameron and Rettger, Jour. Bact., 27, 1934, 86. See Edwards and Bruner, Kentucky FAMILY ENTEROBACTERIACEAE 503 Agr. Exp. Sta. Bull. 400, 1940, 43-70, for a discussion of this species. Rods: 0.5 by 1.0 to 1.5 microns, occur- ring singly. Motile with peritrichous flagella. Gram-negative. Gelatin colonies: Small, circular, gray- ish, granular, becoming yellowish-brown. Gelatin stab: Flat surface growth. No liquefaction. Agar colonies: Small, circular, grayish, entire to undulate. Agar slant: Filiform, grayish, moist, entire growth. Broth: Turbid. Litmus milk: Slightly acid, becoming alkaline. Potato: Grayish -white streak. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, arabinose, maltose, dextrin, mannitol, sorbitol and inositol. Acid from glycerol. No action on lactose, sucrose, raffinose, inulin, salicin or adonitol. Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide produced. Optimum temperature 37 °C. Aerobic, facultative. Antigenic structure: [I], IV, [V],XII: i: 1, 2, 3... Source: Isolated during a mouse ty- phoid epidemic in the Hygienic Institute of Greifswald, German}-. Habitat: Causes food-poisoning in man. A natural pathogen for all warm- blooded animals. This type occurs more frequently than any other type not con- fuied to a specific host. Also found in snakes bj' Hinshaw^ and McNeil (Amer. Jour. Vet. Res., 6, 1945, 264). 4a. iSalmonella typhiinuriuni (Type Binns). (Bacillus paratyphcsus B Binns type, Schtitze, Lancet, 1, 1920, 93; Group VI of Hecht-Johansen, Copenhagen, 1923; Typus-Binns, Kauffmann, Zbl. f . d. ges. Hyg., ^0, 1931, 273; Salmonella iij- phimuriian var. Copenhagen, Kauffmann, Ztschr. f. Hyg., 116, 1934, 368; Salmon- ella typhi-murium var. Binns, Schtitze et al.. Jour. Hyg., 34, 1934, 339; Sal- monella aerirycke var. Storrs, Edwards, Jour. Bact., 30, 1935,471.) Morphology and cultural characters indistinguishable from those of Sal- monella typhimurium, except some strains ferment maltose late or arc anaerogenic. Antigenic structure: [I], IV, XII: i: 1, 2, 3 . . . (Edwards, Jour. Hyg., 36, 1936, 348). Many colonies may be ex- amined before the specific phase flagellar antigen is demonstrated. Differs from Salmonella typhimurium in lacking anti- gen V. Source : Isolated by Dr. McNee from a case of food poisoning in man, France, 1919. Habitat: Natural host the pigeon, and may infect other animals, including man. 5. Sahnonella sp. (Type Koln). {Sal- monella koln Sievers, Cent. f. Bakt., I Abt., Orig., 150, 1943, 52; Salmonella coeln Kauffmann, Acta Path, et Micro- biol. Scand., Suppl. 54, 1944, 33.) Antigenic structure: IV, V, XII: y: 1,2, 3 . . . Source: A single culture isolated from a human case of enteritis. Habitat: Not reported from other sources as yet. 6. Salmonella sp. (Type StanlejO- (Bacillus paratyphcsus B, Stanlej^ type, Schtitze, Lancet, 1, 1920, 93; Salmonella sianleyi Haupt, Ergebnisse d. Hyg., 13, 1932, 673; Salmonella Stanley type. White, Med. Res. Council, Spec. Rept. Ser. 103, 1926, 19; Salmonella Stanley Warren and Scott, Jour. Hyg., 29, 1929, 415; Typus Stanlej', Kauffmann, Ztschr. Hyg., HI, 1930, 210.) Antigenic structure: IV, V, XII: d: 1, 2 . . . Source : Isolated from cases of human food poisoning in Stanley', England by Hutchens (1917). 504 IVIANITAL OF DETERMINATIVE BACTERIOLOGY Habitat: Not known as a natural pathogen of animals. 7. Salmonella sp. (Type Heidelberg). {Bacterium enteriiidis, Typus Heidel- berg, Habs, Cent. f. Bakt., I Abt., Orig., 130, 1933, 367; Salmonella heidelberg Schutze et al., Jour. Hyg., 34, 1934, 340.) Antigenic structure: IV, V, XII: r: 1,2, 3.. . Source: Isolated from cases of human food poisoning in Heidelberg, Germany. Habitat: Not known as a natural pathogen of animals. 8. Salmonella sp. (Type Chester). (Salmonella Chester Kauffmann and Tes- dal, Ztschr. f. Hyg., 120, 1937, 168.) Antigenic structure: IV, [V], XII: e, h: e, n, X . . . Source: Isolated by W. H. Grace, Chester, England, from gastroenteritis in man. Typed by Kauffmann and Tes- dal {loc. cit.). Habitat: Has usually been found in human feces. 9. Salmonella sp. (Type San Diego). {Salmonella san diec/o Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940,429.) Antigenic structure: IV, [V], XII: e, h: e, n, Zn . . . Source: Originally isolated from cul- tures sent to Dr. Kauffmann by Dr. K. F. Meyer who obtained them from an out - break of food poLsoning near San Diego, California. Also reported from Den- mark, Uruguay and Kentucky. Habitat: Usually has been isolated from human feces, but has been found in birds and other animals. 10. Salmonella sp. (Type Salinas). {Salmonella salinatis Edwards and Bruner, Jour. Bact.,U, 1942, 289.) Antigenic structure: IV, XII: d, e, h: d, e, n, Zi5 . . . By cultivation in semi -solid agar con- taining agglutinating serum for Sal- monella typhosa, an organism having the antigenic formula for Salmonella sp. (Type San Diego) was isolated. Source: From rat feces collected by Dr. Henry Welch near Salinas, Cali- fornia. Habitat: Also found in normal human carriers. 11. Salmonella sp. (Type Saint Paul). (Salmonella saint paid Edwards and Bruner, Jour. Inf. Dis., 66, 1940, 220.) Antigenic structure: I, IV, V, XII: e, h: 1, 2, 3 . . . Source: A single culture isolated from the liver of a turkey poult by Dr. B. S. Pomeroy, St. Paul, Minnesota. Two cases in man. Habitat: Also reported from hogs. 11a. Salmonella sp. (Type Zagreb). (Salmonella zagreb Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940, 351.) Antigenic structure: IV, V, XII: e, h: 1,2.... This is a minor tj'pe of No. 11. Source: Culture received by Dr. Kauffmann under the label S. reading from Dr. N. Cernozubov of Zagreb, Jugoslavia. Habitat: Not reported from other sources as yet. 12. Salmonella sp. (Type Reading). (Bacillus paratyphosus B, Reading type, Schutze, Lancet, 1, 1920, 93; Salmonella reading Schutze, Brit. Jour. Exp. Path., 11, 1930, 34; Typus Reading, Kauffmann, Zentbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella readingensis Haupt, Ergeb- nisse d. Hyg., 13, 1932, 673.) Antigenic structure: IV, XII: e, h: 1, 5 Source: Isolated from the Reading, England water supply by Dr. H. Schutze. Also found in hogs (Edwards and Bruner, Jour. Inf. Dis., 72, 1943, 64). Habitat: A cause of gastroenteritis in man. FAMILY ENTEROBA('TERIACEAE 505 12a. Salmonella sp. (Type Kaposvar). (Salmonella kaposvar Rauss, Cent. f. Bakt., I Abt., Orig., U7, 1941, 253; also see Kauffmann, Die Bakteriologie der Salmonella-Gruppe, Kopenhagen, 1941, 212.) Antigenic structure: IV, V, XII: e, (h) 1, 5. . . . This is a minor type of No. 12. Source: From the feces of three mem- bers of a family suffering from gastro- enteritis. Habitat: Not reported from other sources as yet. 13. Salmonella sp. (Tj^pe Kaapstad). (Salmonella reading var. kaapstad Hen- ning, Rhodes and Gordon-.Johnstone, Onderstepoort Jour. Vet. Sci. Animal Ind., 16, 1941, 103; Salmonella kaapstad Kauffmann, Acta Path, et Microbiol. Scand., 19, 194:2, 523.) Antigenic structure: IV, XII: e, h: 1,7.... Source: From a child with meningitis. Habitat: Not known from other sources as yet. 14. Salmonella sp. (Type Derby). (Ba- cillus enteritidis Peckham, Jour. Hyg., 22, 1923, 69; Derby type. Savage and White, Med. Res. Council Spec. Rept. Ser. 91, 1925, 19; Salmonella derby War- ren and Scott, Jour. Hj^g., 29, 1929, 415; Salmonella derbyensis Haupt, Ergebnisse d. Hyg., IS, 1932, 673.) Antigenic structure: [I], IV, XII: f, g:-- Source: Isolated from tank water at Derby, England. Habitat: Widely distributed. Found in human feces, lymph glands of hogs, chickens, etc. 15. Salmonella sp. (Type Essen). (Salmonella essen 173 Hohn and Herr- mann, Cent. f. Bakt., I Abt., Orig., 135, 1936, 505.) Antigenic structure : IV, XII : g, m : — . Source: Isolated from the feces of an infant, Essen, Germany. Habitat : Known only from human sources. 16. Salmonella sp. (Type Budapest). (Salmonella btidapest Rauss, Ztschr. f. Immunitatsf., 95, 1929, 489.) Antigenic structure: I, IV, XII: g, t:— . Source: Originally isolated in Buda- pest from 3 normal persons and from 3 persons with enteric fever. Habitat: Known only from human sources. 17. Salmonella sp. (Tj'pe California). (Salmonella California Edwards, Bruner and Hinshaw, Jour. Inf. Dis., 66, 1940, 127; Hinshaw, Hilgardia, 13, 1941, 583.) Antigenic structure: IV, XII, g, m, t:— . Sotirce: Six cultures isolated from in- fected turkey poults from California. The seventh culture was isolated from a turkey in a second outbreak of the infec- tion. Reported by Pomeroy and Fen- stermacher (Jour. Amer. Vet. Med. Assoc, 94, 1936, 90). Also found in hogs and man (Edwards and Bruner, Jour. Inf. Dis., 72, 1943, 64). Habitat: Also reported from chickens and ducks. Widely distributed. 18. Salmonella sp. (Type Branden- burg). (Typus-Brandenburg, Kauff- mann and ]\Iitsui, Ztschr. f. Hyg., HI, 1930, 740; Kauffmann, Zentbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella branden- burgensis Haupt, Ergebnisse d. Hyg., 13, 1932, 673; Salmonella brandenburg Schtitze et al.. Jour. Hyg., 34, 1934, 540.) Antigenic structure: IV, XII: 1, v: e, n, Zi5. . . . See Kauffmann, Ztschr. f. Hyg., 118, 1936, 540. Source : Isolated from a case of gastro- enteritis at the Virchow Hospital of Berlin. 506 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat: Known only from human sources. 19. Salmonella sp. (Type Bispebjerg). {Salmonella bispebjerg Typus, Kauff- mann, Ztschr. f. Hyg., 118, 1936, 540.) Antigenic structure: I, IV, XII: a: e, n, X. . . . Source: Isolated from a case of gastro- enteritis at the Bispebjerg Hospital in Copenhagen. Habitat: Not reported from other sources as yet. 20. Salmonella abortivoequina (Good and Corbett) Bergey et al. {Bacillus abortivus equinus Good and Corbett, Jour. Inf. Dis., 13, 1913, 53; Bacillus abortus equi Meyer and Boerner, Jour. Med. Res., 29, 1913, 330; Bacillus abor- tivo-equinus Good and Corbett, Jour. Inf. Dis., 18, 1916, 586; Bacillus abortus equinus Weiss and Rice, Jour. Med. Res., 35, 1907, 403; Bacillus abortivus Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 477; Bacillus abortus-equi Holland, Jour. Bact., 5, 1929, 216; Bac- terium abortum-equi Holland, ibid.; Ber- gey et al.. Manual, 1st ed., 1923, 217; Bacillus enteritidis B, Typ. equinus Januschke, Ztschr. f. Infektionskr. d. Haustiere, 29, 1924, 182; Salmonella abortus-equi Bergey et al.. Manual, 2nd ed., 1925, 236.) From Latin, aborting and equine. Antigenic structure: IV, XII: — : e, n, X. . . . Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Source: Isolated from afterbirth of mares that had aborted. Habitat : A natural pathogen of mares, causing abortion. Infectious for guinea pigs, rabbits, goats, cows, producing abortion. 21. Salmonella sp. (Type Arechava- leta). {Salmonella arechavaleta Hor- maeche and Peluffo, quoted from Hor- maeche et al.. Jour. Bact., 47, 1944, 323.) Named in honor of Prof. Arechavaleta of UruguaJ^ Antigenic structure: IV, [V], XII: a: 1,7.... Source : From a human case of gastro- enteritis. Also found by Dr. P. R. Edwards among cultures sent to him from the Canal Zone for identification. Habitat: Known only from human sources. 22. Salmonella abortusovis (Lovell) Schiitze et al. {Bacillus paratyphi abortus ovis Schermer and Ehrlich, Cent, f. Bakt., I Abt., Ref., 73, 1922, 252; Ba- cillus enteritidis C, Typ. ovis Januschke, Ztsch. f. Infektionskr. d. Haustiere, 27, 1924, 182; Bacterium abortus ovis Lovell, Jour. Path, and Bact., 84, 1931, 13; Typus-Abortus ovis, Kauffmann, Zent- Ibl. f. d. ges. Hyg., 25, 1931, 273; Schiitze et al.. Jour. Hyg., 34, 1934, 340.) Antigenic structure: IV, XII: c: 1, 6 Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Source : Isolated from cases of abortion in sheep. Habitat: Not known to infect any other animal. 23. Salmonella sp. (Type Altendorf). {Salmonella altendorf Hohn, Cent. f. Bakt., I Abt., Orig., I46, 1940, 218.) Antigenic structure: IV, XII: c: 1, 7 Source: Isolated from a case of acute gastroenteritis from Altendorf, Ger- many. Habitat: Not reported from other sources as yet. 24. Salmonella sp. (Type Texas). {Salmonella texas Watt, De Capito and Moran, U. S. Public Health Repts., 62, 1947, 80S.) Antigenic structure: IV, V, XII: k: e, n, Z16. . . . Source: Isolated by Dr. James Watt from the feces of a boy convalescing from diarrhoea. FAMILY ENTEROBACTERIACEAE 507 Habitat: Not reported from other sources as yet. 25. Salmonella abortusbovis Kauff- mann. (Kauffmann, Ztschr. f. Hyg., 120, 1937, 194.) Antigenic structure: [I], IV, XXVII, XII: b: e, n, x. . . . Liquefies gelatin (Kauffmann, Ztschr. f. Hyg., 117, 1936, 778). Source: Isolated and incompletely typed by H. Bernard, Ztschr. f. Hyg., 117, 1935, 352. Habitat: Normally found in cattle, causing abortion. Occasionally occurs in man. 26. Salmonella sp. (Tjpe Bredeney). {Salmonella hredeney Kauffmann, Ztschr. f. Hyg., 119, 1937, 356.) Antigenic structure: I, IV, [XXVII], XII: 1, v: 1, 7. . - . Source: Found by Hohn and Herr- mann in Bredeney, Germany. Typed by Kauffmann {loc. cit.). From cases of human gastroenteritis and an abscess of lower jaw. Habitat : Isolated from human sources. Also found in normal hogs and chickens. 27. Salmonella sp. (Type Schleiss- heim). {Salmonella, schleissheim Kauff- mann and Tesdal, Ztschr. f. Hyg., 120, 1937, 171.) Antigenic structure: IV, XXVII, XII: b, Zio: — . Liquefies gelatin (Kauffmann and Tesdal, loc. cit.). Source: Isolated by Hopfengartner (Miinchener tierarz. Wchnschr., 1, 1929, 185) in Schleissheim. From cattle. Typed by Kauffmann and Tesdal {loc. cit.). Also found by Tillmanns in the liver of a horse (Ztschr. f. Fleisch. u. Milch Hyg., 50, 1940, 109). Caused an outbreak of gastroenteritis in 30 persons (Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940, 1). Habitat: Apparently widely dis- tributed. 28. Salmonella sp. (Type Schwarzen- grund) . {Salmonella schwarzengrund Kauffmann, Acta Path, et Microbiol. Scand., Suppl. 44, 1944, 34.) Antigenic structure: I, IV, XXVII, XII: d: 1, 7. Source: A single culture isolated by Dr. J. Hohn from a human case of en- teritis that occurred in Schwarzengrund, near Breslau, Germany. Habitat: Not reported from other sources as yet. 29. Salmonella hirschfeldii Weldin. {Bacillus paratyphosus /Ss Weil, Wien. klin. Wchnschr., 30, 1917, 1061; Bacillus erzinjan Neukirch, Ztschr. f. Hyg., 85, 1918, 103; Paratyphoid C bacillus, Hirschfeld, Lancet, 1, 1919, 296; "Para- C", Mackie and Bowen, Jour. Roy. Army Med. Corps, 33, 1919, 154; Bacillus paratyphosus C Andre wes and Neave, Brit. Jour. Exp. Path., 2, 1921, 157; Paratyphus Ni, Iwaschenzoff, Arch. f. Schiffs- u. Trop. Hyg., 30, 1926, 1 ; Weldin, Iowa Sta. Coll. Jour. Sci., 1, 1927, 161; Bacterium hirschfeldii Weldin, ibid., 161; Typus-Orient, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella paratyphi C Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Salmonella paratyphosus C Castellani and Chalmers, ibid., 952.) Named for Hirschfeld who worked with this organism. Rods: 0.3 to 0.5 by 1.0 to 2.5 microns, occurring singly. Motile with peritri- chous flagella. Gram-negative. Gelatin colonies: Grayish, smooth, flat, glistening, margin irregular. Gelatin stab: Flat, grayish surface growth. No liquefaction. Agar colonies: Grayish, moist, smooth, translucent. Broth: Turbid. Litmus milk: Slightly acid, becoming alkaline. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, maltose, arabinose, xylose, dextrin, trehalose, mannitol, dulcitol and sor- bitol. No action on lactose, sucrose, 508 MANUAL OF DETERMINATIVE BACTERIOLOGY salioin, adonitol or inositol. Rarely may fail to form gas from sugars (Nabih, Jour. Hyg., U, 1941, 39). Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide produced. Optimum temperature 37 °C. Aerobic, facultative. Antigenic structure: VI, VII, [Vi]: c: 1,5. . . . Source: Isolated from cases of enteric fever in man. Habitat: A natural pathogen of man causing enteric fever. 30. Salmonella choleraesuis (Smith) Weldin. (Probably not the Bacillus of swine plague, Klein, Report of the Medi- cal Officer of the Local Gov. Bd., Eng- land, 1877-78, Supplement, p. 168; Bac- terium of swine plague, Salmon, U. S. Dept. Agr. Bur. An. Ind. Ann. Rep., 1885, 212; Bacterium of hog cholera, Salmon, ihid., 1886, 20; Bakterium der Schweinepest, Selander, Cent. f. Bakt., 3, 1888, 361; Pasteurella mlmoni Trevi- san, I generi e le specie delle Batteriacee, 1889, 21; Bacterium, cholerae suis Th. Smith, U. S. Dept. Agr. Bur. An. Ind., Bull. 6, 1894, 9; Swine-feverbacillus, Klein, Cent. f. Bakt., I Abt., 18, 1895, 105; Bacillus suipestifer Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 401; Bacterium cholerae suum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 3, 1896, 233; Bacterium suipesti- fer Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 70; Bacillus cholerae suum Migula, Syst. d. Bakt., 2, 1900, 759; Le microbe du hog-cholera, Lig- nieres. Bull. Soc. Cent. M^d. Vet., see Rec. de m^d. v^t., Paris, S^r. 8, 7, 1900, 389; Bacillus salmoni Chester, Manual Determ. Bact., 1901, 210; Bacterium intestinale suis Le Blaye and Guggen- heim, Manuel Pratique de Diagnostic Bacteriologique, 1914; Bacillus suis Krumwiede, Kohn and Valentine, Jour. Med. Res., 88, 1918, 89; Bacterium (Salmonella) cholera suis Buchanan, Jour. Bact., 3, 1918, 53; Salmonella suipestifer Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Bacillus cholerae-suis Winslow, Kligler and Rothberg, Jour. Bact., 4, 1919, 476; Bacterium cholerae-suis Holland, Jour. Bact., 5, 1920, 217; Bacillus paratypho- sus B (Arkansas type), Schtitze, Lancet, 2, 1920, 93; included in Group I suipesti- fer, Andrewes and Neave, Brit. Jour. Exp. Path., 2, 1921, 157; Weldin, Iowa Sta. Coll. Jour. Sci., 1, 1927, 155; Ty- pus suipestifer Amerika, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; the Ameri- can Salmonella suipestifer of many authors.) From Latin, hog cholera. Salmonella choleraesuis (Smith) Wel- din is the type species of the genus Sal- monella. Rods: 0.6 to 0.7 by 2.0 to 3.0 microns, occurring singly. Motile with four to five peritrichous flagella. Gram-nega- tive. Gelatin colonies: Grayish, smooth, fiat, glistening; margin irregular. Gelatin stab: Flat, grayish surface growth. No liquefaction. Agar colonies: Grayish, moist, smooth, translucent. Agar slant: Grayish, moist, smooth, translucent growth. Broth: Turbid, with thin pellicle and grayish-white sediment. Litmus milk: Slightly acid, becoming alkaline, opalescent, translucent to yel- lowish-gray. Potato: Grayish -white streak becom- ing brownish. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, xylose, maltose, glycerol, mannitol, dulcitol, rhamnose, sorbitol and dextrin. Arabinose, inosi- tol, lactose, sucrose, salicin, inulin, raifinose and trehalose not attacked. Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide not produced. Optimum temperature 37 °C. Aerobic, facultative. Antigenic structure: VI, VII: c: 1, FAMILY EXTEROBACTERIACEAE 509 5 . . . Serologically identical with Sal- monella typhisuis, and cross-aggluti- nates to a varying degree with a number of other serotypes. Habitat and source: Natural host the pig as an important secondary invader in the virus disease, hog cholera. Does not occur as a natural pathogen in other animals, although lethal for mice and rabbits on subcutaneous injection. Oc- casionally gives rise to acute gastro- enteritis and enteric fever in man. 30a. Salmonella choleraesuis var. Knn- zendorf Schiitze et al. The synonyms up to and including Weldin, 1927 for Salmonella choleraesuis apply equally well to the var. Kunzen- dorf, for these were not separated with certainty until 1926 (White, Med. Res. Council, London, Spec. Rep. Ser. 103, 27). Re-examined serologically a num- ber of previously described strains agree with this variety. (Paratyphus C Ba- cillus, Heimann, Cent. f. Bakt., I Abt., Orig., 66, 1912, 211; Paratyphosus C, Weil and Saxl, Wien. klin. Wchnschr., 30, 1917, 519; Typus-suipestifer Kun- zendorf, Pfeiler, Ztschr. f. Infektskr. d. Haust., 20, 1920, 218; Bacillus para- typhosus B, G. type, Schiitze, Lancet, 1, 1920, 93; Bacillus paratyphosus C, Dud- geon and Urquart, Lancet, 2, 1920, 15; included in Group II suipestifer, An- drewes and Xeave, Brit. Jour. Exp. Path., 2, 1921, 157 and Group V of Hecht- Johansen, Copenhagen, 1923; Salmonella suipestifer (European variety) Schiitze. Brit. Jour. Exp. Path., 11, 1930 34; Typus-suipestifer Kunzendorf, Kauff- mann, Zbl. f. d. gen. Hyg., 25, 1931, 273; Salmonella choleraesuis var. kunzen- dorf Schiitze et al., Jour. Hyg., 34, 1934, 341; the European Salmonella suipestifer of many authors.) Indistinguishable from Salmonella choleraesuis in morphology and cultural characters, except that the Kunzendorf variety forms hydrogen sulfide. Antigenic structure: VI, VII: [c]: 1, 5. Differs from Salmonella choleraesuis in lacking the specific flagellar phase; serologically identical with Salmonella typhisuis var. voldagsen. Source: From pigs with swine fever and once from a monkey in captivity. Habitat: Causes acute gastro enteritis and enteric fever in man. Also found in cattle, sheep, carnivora and chickens. 31. Salmonella typhisuis (Glasser) Schiitze et al. (Bacillus typhisuis Glas- ser, Deutsche tierarztl. Wchnschr., 17, 1909, 513; included in the Ferkeltyphus bacilli of German literature, Dammann and Stedefeder, Arch. f. wiss. u. prakt. Tierheilk., 36, 1910, 432; Bacillus glasser Neukirch, Ztschr. f. Hyg., 85, 1918, 103; Bacterium typhisuis Holland, Jour, Bact., 5, 1920, 221; included in Group I suipestifer, Andrewes and Neave, Brit. Jour. Exp. Path., 2, 1921, 157; Typus-Glasser, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Schutze et al.. Jour. Hyg., 34, 1934, 342.) From Greek, typhus and Latin, pig. Rods: 0.6 to 0.7 by 2.0 to 3.0 microns, occurring singly. Motile with four to five peritrichous flagella. Gram -nega- tive. Gelatin colonies: Grayish, smooth, flat, glistening, edge entire. No lique- faction. Agar colonies: Grayish, moist, smooth, translucent. Broth: Turbid. Litmus milk: Slightly acid or neutral. Indole not formed. Nitrites produced from nitrates. Forms gas slowly and sparsely from all substances. Growth poor on all or- dinary media. Acid from arabinose, xylose and tre- halose. Delayed or variable fermenta- tion from dextrin, maltose, rhamnose, dulcitol, sorbitol. Mannitol not fer- mented or very slowly. Inositol not fermented. No HoS produced. Optimum temperature 37°C. Aerobic, facultative. Antigenic structure: Identical with Salmonella choleraesuis, from which the 510 MANUAL OF DETERMINATIVE BACTERIOLOGY organism differs in respect to arabinose and trehalose. Antigenic structure VI, VII: c: 1, 5 Habitat: Infects only the pig. 31a. Salmonella typhisuis var. vol- dagsen Schiitze et al. (Included in Ferkeltyphus bacilli, Dammann and Stedefeder, Arch. f. wiss. u. prakt. Tierheilk., 36, 1910, 432; Bacillus vol- dagsen Neukirch, Ztschr. f. Hyg., 85, 1918, 103; included in Group II suipesti- fer, Andrewes and Neave, Brit. Jour. Exp. Path., 2, 1921, 157; Typus-voldag- sen, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella typhisuis var. voldagsen, Schiitze et al., Jour. Hyg., 34, 1934, 342.) Morphology and cultural characters identical witli those of Salmonella typhisuis. Antigenic structure: VI, VII: [c]: 1, 5. . . . Identical with that of Salmonella choleraesuis var. Kunzendorf ivom which. species the organism differs culturally. Habitat: Infects only the pig. 32. Salmonella sp. (Type Thompson). (Thompson type of Salmonella, Scott, Jour. Hyg., 25, 1925, 398; Typus-Thomp- son-Berlin, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella thompsoti Schutze et al., Jour. Hyg., 84, 1934, 343.) Named after the family involved in the outbreak. Antigenic structure, VI, VII; k: 1, 5. . . . Source: Isolated from food poisoning in man. Also found in chickens and turkeys (Edwards and Bruner, Jour. Inf. Dis., 72, 1943, 64). Habitat: Widely distributed in warm- blooded animals. 32a. Salmonella sp. (Type Berlin) (Type Thompson). (Typus-Berlin, Kauffmann, Cent. f. Bakt., I Abt., Ref., 94, 1929, 282; Typus C Berlin, Boecker and Kauffmann, Cent. f. Bakt., I Abt., Orig., 116, 1930, 458; Typus-Thompson- Berlin, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella thompson var. berlin Schutze et al.. Jour. Hyg., 34, 1934, 343.) Antigenic structure: VI, VII: [k] : 1, 5 Source: Isolated from food poisoning in man. Not known to be a natural pathogen of animals. Habitat: A natural pathogen of man causing food poisoning. 33. Salmonella sp. (Type Montevideo). (Salmonella montevideo Hormaeche and I'eluffo, Arch. Urug. de Med., Cirug. y Espec, 9, 1936, 673.) Antigenic structure: VI, VII: g, m, s: — . Source: Originally isolated from hu- man sources in Montevideo from an ape that died of an enterocolitis, and mesen- teric glands of healthy hogs; also re- ported from chickens and powdered eggs (Schneider, Food Research, 11, 1946, 313). Habitat: Apparently widely dis- tributed. 34. Salmonella sp. (Type Oranien- burg). (Typus-Oranienburg, Kauff- mann, Ztschr. f. Hyg., Ill, 1930, 223; Salmonella oranienhurgensis Haupt, Ergebnisse der Hyg., 13, 1932, 673; Salmonella oranienburg Schutze et al., Jour. Hyg., 34, 1934, 343.) Antigenic structure: VI, VII: m, t: — . Source: From the feces of a child in a children's home near Oranienburg. La- ter isolated from gastroenteritis in man. Also from cjuail, chickens and powdered eggs (Schneider, loc. cit.). Habitat: Reported from human sources, from hogs and from birds. 35. Salmonella sp. (Type Virchow). (Typus-Virchow, Kauffmann, Ztschr. f. Hyg., Ill, 1930, 221; Salmonella vir- chowii Haupt, Ergebnisse der Hyg., 13, 1932, 673; Salmonella virchow Schutze et al., Jour. Hyg., 34, 1934, 343.) Antigenic structure: VI, VII: r: 1, 2,3.... FAMILY ENTEROBACTERIACEAE 511 Source: Isolated from food poisoning in a man at the Rudolf Virchow Hospital in Berlin. Habitat: A natural pathogen of man causing food poisoning. 36. Salmonella sp. (Type Oslo). {Sal- monella Oslo Tesdal, Ztschr. f. Hyg., 11.9, 1937,451.) Antigenic structure: VI, VII: a: e, n, X. Source: Isolated in Oslo, Norway from cases of gastroenteritis in man. Habitat: Not reported from other sources as yet. 37. Salmonella sp. (Type Amersfoort). (Salmonella amersfoort Henning, Jour. Hyg., 37, 1937,561.) Antigenic structure: VI, VII: d: e, n, X. . . . Source: Originally isolated from chickens from Amersfoort, Transvaal. Later found in a human mixed infection with Salmonella typhi murium. Habitat: Not reported from other sources as yet. 38. Salmonella sp. (Type Braenderup). {Salmonella braenderup Kauffmann and Henningsen, Ztschr. f. Hyg., 120, 1937, 640.) Antigenic structure: VI, VII: c, h: e, n, Zi5. . . . Source : Isolated from a case of human gastroenteritis in Braenderup, Denmark. Also from a cat in the same home that had died from a diarrhoea. Reported later from So. Africa (see Kauffmann, Die Bakteriologie der Salmonella - Gruppe, Kopenhagen, 1941, 2.37). Habitat: Apparentl}' wide]}' dis- tributed. 39. Salmonella sp. (Type Potsdam). (Typus-Potsdam, Kauffmann and Mitsui, Ztschr. f. Hyg., Ill, 1930, 740; Salmonella potsdamensis Haupt, Ergeb- nisse der Hyg., 13, 1932, 673; Salmonella potsdam Schiitze et al., Jour. Hyg., 34, 1934, 343.) Antigenic structure: VI, VII: 1, v: e, n. zi5. . . . Source: Isolated from food poisoning in man at Potsdam, Germany. Habitat: A natural pathogen of man causing food poisoning. 40. Salmonella sp. (Type Bareilly). {Salmonella, Type Bareilly, Bridges and Scott, Jour. Roy. Army Med. Corps, 56, 1931, 241; Salmonella bareilly Schiitze et al.. Jour. Hyg., 34, 19.34, 343.) Antigenic structure: VI, VII: y: 1, 5 Source: Isolated in 1928 from cases of mild enteric fever that occurred in Bareilly, India. Also reported from chickens (Kauffmann, Die Bakteriologie der Salmonella -Gruppe, Kopenhagen, 1942, 235). Habitat: A natural pathogen of man causing gastroenteritis and enteric fever. Widely distributed in fowls. 41. Salmonella sp. (Type Hartford). {Sahnonella hartford Edwards and Bruner, Jour. Inf. Dis., 69, 1941, 223.) Antigenic structure: VI, VII: }'•: c, n, X. . . . Source: One culture isolated from the stool of a man with persistent diarrhoea by Dr. E. K. Borman, Hartford, Conn. Habitat: Not reported from other sources as }^et. 42. Salmonella sp. (Type Mikawa- sima*). (Salmonella bareilly var. mika- wasima Hatta, Japan Jour. Exper. Med., 16, 1938, 201; Salmonella mikawa- sima Hormaeche, quoted from Schiitze et al., Proc. 3rd Internat. Cong. Mi- crobiol., 1940, 337; also see Kauffmann, Acta Path, et Microbiol. Scand., 16, 1939, 347 and ibid., 17, 1940, 429.) Antigenic structures: VI, VII: y: e, n, zio. . . . * Correct spelling according to Prof. Kojima. 512 MANUAL OF DETERMINATIVE BACTERIOLOGY Source: Isolated from a rat by Prof. Kojima and Prof. Hatta, 1937. Habitat: Not reported from other sources as j'et. 43. Salmonella sp. (Type Tennessee). (Salmonella tennessee Bruner and Ed- wards, Proc. Soc. Exp. Biol, and Med., SO, 1942, 174.) Antigenic structure: VI, VII: Z29: — ■ Source : Culture isolated from feces of normal carrier by Dr. W. C. Williams, State Dept. of Health, Nashville, Tennessee. Habitat: Also reported from turkej^s and powdered eggs. 44. Salmonella sp. (Type Concord). (Salmonella var. concord Edwards and Hughes, Jour. Bact., 47, 1944, 574.) Antigenic structure: VI, VII: 1, v: 1,2, 3. . . . Source: Two cultures isolated by Dr. J. R. Beach and one by Dr. C. U. Duck- worth from fatal infections in chicks (U. tS. A.) and one by Dr. Joan Taylor from the stool of a person affected with gastroenteritis (England). Habitat: Also reported from turke3'S. 45. Salmonella sp. (Type Infantis). (Salmonella infantis Wheeler and Bor- man. Jour. Bact., 46, 1943, 481.) Antigenic structure: VI, VII: r: 1, 5. . . . Source: Isolated at Hartford, Con- necticut from the blood of an infant. Subsequently also from stools. Habitat: Not reported from other sources as yet. 46. Salmonella sp. (Type Georgia). (Salmonella georgia Morris, Brim and Sellers, Amer. Jour. Pub. Health, 34, 1944, 1279; Seligmann, Saphra and Was- sermann, Amer. Jour. Hyg., 40, 1944, 227.) Antigenic structure: VI, VII: b: e, n, Zj5. . . . Source: Isolated by Miss Jane Morris from the feces of a 16-year-old boy dur- ing routine examination of food handlers, State Dept. of Health, Atlanta, Georgia. Habitat: Not reported from other sources as yet. 47. Salmonella sp. (Type Papua). (Salmonella papnana Wilcox, Edwards and Coates, Jour. Bact., 49, 1945, 514.) Antigenic structure: VI, VII: r: e, n, Z]6. . . . Source: Isolated by Lt. Goldwasser from human feces from Port Moresby in Papua, New Guinea. Habitat: Not reported from other sources as yet. 48. Salmonella sp. (Type Richmond). (Salmonella richmond Moran and Ed- wards, Proc. Soc. Exp. Biol, and Med., 62, 1946, 294.) Antigenic structure: VI, VII: y: 1, 2, 3. . . . Source : Isolated bj^ Mr. Forest Spindle in Richmond, Virginia from the feces of a child affected with gastroenteritis. Habitat: Isolated as yet from human sources only. 49. Salmonella sp. (Type Cardiff). (Salmonella Cardiff Taylor, Edward and Edwards, Brit. Med. Jour., 1945, i, 368.) Antigenic structure: VI, VII: k: 1, 10. . . . Source: Isolated from human case of gastroenteritis from Cardiff, Wales. Habitat: Isolated as yet from human sources only. 50. Salmonella sp. (Type Daytona). (Salmonella daytona Moran and Ed- wards, Proc. Soc. Exp. Biol, and Med., 62, 1946, 294.) Antigenic structure: VI, VII: k: 1, 6. . . . Source: Isolated by Mrs. Mildred Gal- ton from human feces from Daytona, Florida. Habitat: Not known from other sources as yet. FAMILY ENTEROBACTERIACEAE 513 51. Salmonella sp. (Type Newport). (Paratj-phus /3o, Weil and Saxl, Wien. klin. Wchnschr., 30, 1917, 519; Bacillus paratyphosus B, Newport type, Schiitze, Lancet, 7, 1920, 93; Paratyphus Newport Bacillus, Kauffmann, Cent. f. Bakt., I Abt., Ref., 94, 1929, 282; Salmonella newport Schtitze, Brit. Jour. Exp. Path., 11, 1930, 34; Salmonella newportensis Haupt, Ergebnisse der Hyg., 13, 1932, 673.) Antigenic structure: VI, VIII: e, h: 1,2, 3. . . . Source: Isolated from food poisoning in man, Newport, England. Habitat: Widely distributed in man, cattle, hogs, chickens, etc. Also in snakes (Hinshaw and McNeil. Amer. Jour. Vet. Res., 6, 1945. 264). 51a. Sahnonella sp. (Type Puerto Rico) Kauf!'mann. (Jordan, Amer. Jour. Trop. Dis., U, 1934, 27; Kauff- mann, Cent. f. Bakt., I Abt., Orig., 13^, 1934, 162; Schiitze et al.. Jour. Hyg., 34, 1934, 344.) Antigenic structure: VI, VIII: [e, h] : 1,2,3.... This is regarded as a non-specific variant of Salmonella sp. (Type New- port) by Schiitze et al. (Proc. 3rd In- ternat. Cong. Microbiol., New York, 1940,833). 52. Salmonella sp. (Type Pueris). {Salmonella pueris Wheeler and Borman , Jour. Bact., 46, 1943, 481.) Antigenic structure: VI, VIII: e, h: 1,2.... Source: Isolated at Hartford, Con- necticut from anal swabbings of a 14- year-old boy during an attack of gastro- enteritis complicating measles. Habitat: Not reported from other sources as yet. 53. Salmonella sp. (Type Kottbus). {Salmonella newport var. kotibus Kauff- mann, Cent. f. Bakt., I Abt., Orig., 132, 1934, 162; Salmonella kottbus Schiitze et al., Proc. 3rd Internat. ^licrobiol. Cong., New York, 1940, 834.) Antigenic structure: VI, VIII: e, h: 1, 5. . . . Source : From an acute case of gastro- enteritis in Kottbus, Denmark. Habitat: Not reported from other sources as yet . 54. Salmonella sp. (Type Muenchen). (Typus Miinchen, iNIandelbaum, Cent, f. Bakt., I Abt., Ref., 105, 1932, 377; Salmonella muenchen Schiitze et al.. Jour. Hyg., 34, 1934, 344.) Antigenic structure: VI, VIII: d: 1,2.... Source: Isolated from a fatal case of enteric fever. Habitat: Widelj^ distributed. Re- ported from man, rabbits, hogs, camels and chickens (Kauffmann, Die Bak- teriologie der Salmonella Gruppe, 1941, 244). 54a. Salmonella sp. (Tj'pe Oregon). {Salmonella Oregon Edwards and Bruner, Amer. Jour. Hyg., 34, 1941, 21.) Antigenic structure: VI, VIII: d: 1, 2, 3. . . . Source: Six cultures, one isolated from a turkey by Dr. E. M. Dickinson and five from the mesenteric glands of apparently normal hogs by Dr. H. L. Rubin. This is a minor type of No. 54. Habitat: Also reported from reptiles, chickens and man. Also powdered eggs. 55. Salmonella sp. (Type Manhattan). {Salmonella manhattan Edwards and Bruner, Amer. Jour. Hyg., 34, 1941, 21.) Antigenic structure: VI, VIII: d:, 1,5. .. . Source: Two cultures, one isolated from a chicken by Dr. L. D. Bushnell, Manhattan, Kansas, and the other from a turkey by Dr. W. R. Hinshaw. Also 514 MANUAL OF DETERMINATIVE BACMRIOLOGY from reptiles, hogs and human sources (Edwards and Bruner, Jour. Inf. Dis., 7^, 1942,64). Habitat: Apparently widely dis- tributed. 56. Salmonella sp. (Type Litchfield). (Salmonella Utchfield Edwards and Bru- ner, Jour. Inf. Dis., 66, 1940, 220.) Antigenic structure: VI, VIII: 1, v: 1,2,3. .. . Source: Isolated from the liver of a young turkey poult from Litchfield, Minnesota by Dr. B. S. Pomeroy. .A.lso isolated from a case of food poisoning in man by Miss Georgia Cooper. Habitat : Not reported from any other source, as yet. 57. Salmonella morbificans (Migula) Haupt. (Bacillus bovis 7norbificans Base - nau. Arch, f . Hyg., 20, 1894, 257 ; Bacillus morbificans bovis Kruse, in Flligge, Die Mikroorganismen, 3 Aufl., 2, 1896, 380; Bacterium morbificans bovis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 70; Bacillus morbificans Migula, Syst. d. Bakt., 2, 1900, 747; Flavobac- terium morbificans Bergey et al.. Manual, 3rd ed., 1930, 147; Haupt, Ergebnisse der Hyg., 13, 1930, 673; Salmonella bovis-morbificans Schiitze et al.. Jour. Hyg., S^, 1934, 344.) Antigenic structure: VI, VIII: r: 1, 5. . . . Source: Originally isolated from a septicemia in a cow. Habitat: Also found in rabbits and in gastroenteritis in man. 58. Salmonella sp. (Type Narashino). (Salmonella narashino Nakaguro and Yamashita, quoted from Kauffmann, Die Bakteriologie der Salmonella- Gruppe, Kopenhagen, 1941, 246.) Antigenic structure: VI, VIII: a: e, n, X. . . . Source: From the blood and feces of a person suffering from enteric fever. Found in Japan. Habitat: Not reported from other sources as yet. 59. Salmonellasp. (Type Buenos Aires). (Salmonella bonariensis Monteverde, Na- ture, H9, 1942, 472.) Antigenic structure: VI, VIII: i: e, n, X. . . . Source: Isolated by Dr. Monteverde, Buenos Aires from a mesenteric gland of a normal hog. Habitat: Also reported from normal human carriers and from cases of gastro- enteritis. 60. Salmonella sp. (Type Glostrup). (Salmonella glostrup Kauffmann and Henningsen, Acta Path, et Microb. Scand., 16, 1939, 99.) Antigenic structure: VI, VIII: zio: e, n, zi6. . . . Source: Isolated from cases of gastro- enteritis in a family in Denmark. Also affected their dog. Later isolated in Jugoslavia and in Palestine. Habitat : Evidently widely distributed. 61. Salmonellasp. (Type Duesseldorf). (Salmonella duesseldorf Hohn, Cent. f. Bakt., I Abt., Orig., ^6, 1940, 218.) Antigenic structure: VI, VIII: Z4, zu: Source: Isolated from two patients, one of whom died. Found in Duessel- dorf, Germany. Habitat: Not reported from other sources as yet. 62. Salmonellasp. (Type Tallahassee) . (Salmonella tallahassee Moran and Ed- wards, Proc. Soc. Exp. Biol, and Med., 62, 1946, 294.) Antigenic structure: VI, VIII: z^, Z32: — . Source: Isolated by Mrs. Mildred Gal- FAMILY ENTEROBACTERIACEAE 515 ton from feces of gastroenteritis patients and from normal human carriers, Talla- hassee, Florida. Habitat: Not known from other sources. 63. Salmonella sp. (Type Gatun). (Salmonella gatuni Wilcox and Coates, Jour. Bact., 51, 1946, 561.) Antigenic structure: VI, VIII: b: e, n, X. . . . Source: Isolated from human feces from Gatun, Canal Zone. Habitat: Not known from other sources as yet. 64. Salmonella sp. (Type Amherst). {Salmonella amherstiana Edwards and Bruner, Jour. Immunol., U, 1942, 319.) Antigenic structure: (VIII): 1, v: 1, 6 Source: Isolated by Dr. H. Van Roekel from one of a group of poults affected with a fatal disease. Habitat: Not reported from other sources as j^et. 65. Salmonella sp. (Type Virginia). (Salmonella Virginia Saphra and Selig- mann, Proc. Soc. Exper. Biol, and Med., 58, 1945, 50.) Antigenic structure: (VIII): d: — . Source: Isolated by F. Spindle, Rich- mond, Virginia from the feces of an adult person suffering from a diarrhoea. Habitat: Not known from other sources as yet. 66. Salmonella typhosa (Zopf) White. (Bacillus lies Abdominal-Typhus, Eberth, Arch. f. path. Anat., 81, 1880 and 83, 1881; Typhus bacillen, Gaffky, Mitteil. a. d. kaiserl. Gesundheitsamte, 2, 1884, 372; Bacillus typhosus Zopf, Die Spaltpilze, 3 Aufl., 1885, 126; not Ba- cillus typhosus Klebs, Handbuch d. path. Anat., 1880; Bacilbis typhi Schroeter, in Cohn, Kryptogamen Flora v. Schlesien, 3, 1886, 165; Bacillus typhi abdominalis Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 198; Vibrio typhosus Trevisan, I generi e le specie delle Batteriacee, 1889, 23; Bacterium typhi Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 73; Bacterium typhosum Twort, Proc. Royal Soc, London, 79, B, 1907, 329; Acystia typhi Enderlein, Sitzber. Gesell. Naturf. Freunde, Berlin, 1917, 517; Bacterium (Eberthella) typhi Buchanan, Jour. Bact., 3, 1918, 53; Eberthus typhosus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 936; Eberthella typhi Bergey et al.. Manual, 1st ed., 1923, 223; Eberthella typhosa Weldin, Iowa State College Jour. Sci., 1, 1927, 170; Salmonella typhi Warren and Scott, Jour. Hyg., 29, 1930, 416; White, Jour. Hyg., ^9, 1930, 443.) Named from the disease, tj'phoid fever. The species name typhosa should be used for the typhoid organism when it is placed in any genus other than Ba- cillus in spite of the earlier use of this species name by Klebs for a different organism. There are two reasons for this: (a) This appears to be the proper course to follow under International Rules of Nomenclature (See Art. 54, p. 54) and (b) there is less chance for con- fusion regarding the nature of this or- ganism among English-speaking persons who may carelessly interpret typhi as the name of a typhus rather than a typhoid bacillus. Rods: 0.6 to 0.7 by 2.0 to 3.0 microns, occurring singly, in pairs, occasionally short chains. Motile with peritrichous flagella. Gram-negative. Gelatin colonies: Grayish, transparent to opacjue, with leaf-like surface markings. Gelatin stab: Thin, white, opalescent growth. No liquefaction. Agar colonies: Grayish, transparent to opaque. Agar slant: Whitish-gray, glistening, echinulate, entire to undulate growth 516 MANUAL OF DETERMINATIVE BACTERIOLOGY Broth: Turbid, moderate sediment and delicate pellicle in old cultures. Litmus milk: Slight, transient acidity, followed by a return to neutral or to slight alkalinit}^ Potato: Delicate, moist, slightly spreading, barely visible growth. Acid but no gas from glucose, fructose, galactose, xylose, maltose, raffinose, dextrin, glycerol, mannitol and sorbitol. No action on lactose, sucrose, inulin, rhamnose, inositol, salicin and usually arabinose and dulcitol. Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Indole not formed. No characteristic odor. Nitrites produced from nitrates. Hydrogen sulfide produced. Aerobic, facultative. Optimum temperature 37°C. Antigenic structure: IX, XII, [Vi]: d : — . The somatic antigens are related to those of Salmonella enteritidis and a number of other species of Salmonella. V and W forms are present (Felix and Pitt, Jour. Path, and Bact., 38, 1934, 409; Craigie and Brandon, Jour. Path, and Bact., 43, 1936, 233 and 239). Craigie and Yen (Canadian Public Health Journal. 29, 1938, 448 and 484) by the action of selected Vi phages recog- nize eleven distinct stable types of Salmonella typhosa which have been found to be of epidemiological impor- tance. Source: From the human intestine. Habitat: The cause of typhoid fever. Pathogenic for laboratory animals on parenteral injection. Isolated once from a chicken by Henning, Onderste- poort, So. Africa. Note : This species has previously been regarded as the type species of the genus Eberthella Buchanan (Acystia Enderlein, Sitzber. Gesell. Naturf. Freunde, Berlin, 1917, 317; Buchanan, Jour. Bact., 3, 1918, 53; Eberthus Cas- tellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 934; Lankoides Castellani and Chalmers, ibid., 938; Wesenbergus Castellani and Chalmers, ibid., 940.) 67. Salmonella enteritidis (Gaertner) Castellani and Chalmers. {Bacillus en- teritidis Gaertner, Correspond, d. AUge- mein. Artzl. Verein Thuringen, 17, 1888, 573; Klebsiella enteritidis De Toni and Trevisan, in Saccardo, Sylloge Fun- gorum, 8, 1889, 923; Bacterium enteritidis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 68; Bacillus gaertner Morgan, Brit. Med. Jour., 1, 1905, 1257; Castellani and Chalmers, Manual Trop. Med., 3rd ed., 1919, 930.) Named for the disease, enteritis. Rods: 0.6 to 0.7 by 2.0 to 3.0 microns, occurring singly, in pairs and occasion- ally in short chains. Motile with peri- trichous flagella. Gram-negative. Gelatin colonies: Circular, gray, trans- lucent, granular, entire. Gelatin stab: Abundant surface growth. No liquefaction. Agar colonies: Circular, gray, trans- lucent, moist, smooth, entire. Desko- witz and Buchbinder (Jour. Bact., 29, 1935, 294) describe a variant that pro- duces a soluble yellow pigment where certain peptone is present in the agar. Antigenic structure not determined. Agar slant: Grayish-white, opalescent, smooth, moist, undulate growth. Broth: Turbid, with thin pellicle and grayish-white sediment. Litmus milk: Slightly acid, becoming alkaline, opalescent, translucent to yel- lowish-gray. Potato: Al)undant, moist, yellowish- brown to brown growth. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, arabinose, xylose, maltose, trehalose, dextrin, glycerol, mannitol, dulcitol and sorbitol. No acid or gas from lactose, sucrose, inulin, salicin, raffinose, adonitol and inositol. FAMILY ENTEEOBACTERIACEAE il7 Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide produced. No characteristic odor. Aerobic, facultative. Optimum temperature 37 °C. Antigenic structure: [I], IX, XII: g, m:— . Source: First isolated from feces in an epidemic of meat poisoning at Franken- hausen, Germany. Habitat : Widely distributed, occurring in man. Also in domestic and wild animals, particularly rodents. 67a. Sahnonella enterilidis var. Danysz. (Bahr. Deutsche tierarztl. Wchnschr., 1928, 786 and 1930, 14.5; Typus-Gartner Ratin, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Salmonella enterilidis var. danysz, Schiitze et al.. Jour. Hyg., 3^, 1934, 345; Salmonella danysii Gay et al.. Agents of Disease and Host Resistance, 1935, 650.) Differs from Salmonella enterilidis only in its negative action on gh'cerol in Stern's medium. Source: Isolated by Danysz in 1900. Habitat: A natural pathogen of ro- dents and man. 67b. Salmonella enterilidis var. Chaco. (Sa vino and Menendez, Rev. Inst. Bact., 6, 1934, 347; Kauffmann, Ztschr. f. Hyg., //7, 1935, 401.) Differs from Salmonella enterilidis in its action on dulcitol when tested by the method of Bitter, Weigmann and Habs (Munch, med. Wchnschr., 73, 1926, 940.) Habitat and source: Isolated from cases of fever during the Chaco war, South America. 67c. Salmonella enteritidis var. Essen. (Hohn and Herrmann, Cent. f. Bakt., I Abt., Orig., 133, 1935, 183; ibid., 134, 1935, 277; Kauffmann, Ztschr. f. Hyg., 117, 1935,401.) Differs from Salmonella enteritidis when tested by the method of Bitter, Weigmann and Habs (Miinch. med. Wchnschr., 73, 1926, 940), giving a nega- tive reaction with arabinose and dulcitol. Habitat and source: Isolated from human gastroenteritis, ducks and duck eggs. Note: Jansen (Cent. f. Bakt., I Abt., Orig., 135, 1935, 421) states that the organism named by him Salmonella enteritidis var. Mulheim is in reality Salmonella enteritidis var. Essen. 67d. Salmonella enteritidis var. Jena. (Fournier, Rev. Immunolog., Paris, 6, 1940-41, 264.) Source: Isolated from purulent pleural fluid. Habitat: X^ot reported from other sources as yet. 68. Salmonella sp. (Type Dublin). (Bacillus enteritidis Pesch, Cent. f. Bakt., I Abt., Orig., 98, 1926, 22; Dublin Type, White, Med. Res. Counc, Syst. of Bact., 4, 1929, 86 and White, Jour. Hyg., 29, 1930, 443; Salmonella dublin Warren and Scott, Jour. Hyg., 29, 1930, 415; Tj'pus-Dublin-Kiel, Kauffmann, Zbl. f. d. ges. Hyg., 25, 1931, 273; Sal- monella enteritidis var. dublin Schiitze et al.. Jour. Hyg., 34, 1934, 345.) Antigenic structure: I, IX, XII: g, P:— • Source: From meningitis in children (Pesch, loc. cit.). Also isolated by Dr. J. W. Bigger in Dublin, Eire from a fatal fever following a kidney operation. Typed by Dr. Bruce White {loc. cit.). Habitat: Found in man. A natural pathogen of cattle. Widely distributed in cattle and foxes. Two special fermentative types belong here: (1) Salmonella dublin 2 = Sal- monella dublin var. accra Kauffmann, (2) Salmonella dublin 3 = Sahnonella dublin var. koeln Kauffmann (Die Bak- teriologie der Salmonella-Gruppe, Kop- enhagen, 1941, 252). 69. Salmonella sp. (Type Rostock). (Gartner-Poppe Typus, Bahr, Dtsch. Tierarzt. Wchnschr., 1930, 145; Typus 518 MANUAL OF DETERMINATIVE BACTERIOLOGY Gartner-Rostock, Kauffmann, Ztschr. f. Hyg., Ill, 1930, 221 ; Salmonella enteri- tidis var. rostock Schutze et al., 34, 1934, 345; Salmonella rostockensis Haupt, Ergebnisse der Hyg., IS, 1932, 673.) Antigenic structure: I, IX, XII: g, p, u:— . Source : Originally isolated from cattle by Dr. Poppe in Rostock, Germany. Habitat: N^ot known to infect man. 70. Salmonella sp. (Type Moscow). (Paratypus Ci, Weigmann, Cent. f. Bakt., I Abt., Grig., 97, 1925, Beiheft, 299; Salmonella Type Moscow, Hicks, Jour. Hyg., 29, 1929, 446; Salmonella nioscow Warren and Scott, Jour. Hyg., 29, 1929, 446; Typus Gartner-Moskow, Kauffmann, Ztschr. f. Hyg., Ill, 1930, 229; Salmonella moscoivaensis Haupt, Ergebnisse der Hyg., 13, 1932, 673; Sal- monella enieritidis var. moscow, Schutze et al., 34, 1934, 345). Antigenic structure : IX, XII : g, q : — . Source: From patients with enteric fever. Isolated in Moscow, Russia. Habitat: Infects man, horses, cattle. 71. Salmonella sp. (Type Blegdam). {Salmonella blegdam Kauffmann, Ztschr. f. Hyg., 117, 1935,431.) Antigenic structure: IX, XII: g, m, q:— . • Source: Isolated in 1929, at State Serum Institute, Copenhagen from the blood of a pneumonia patient. Also found in the blood of a patient by Dr. Fournier, in Shanghai, China (Kauff- mann, Die Bakteriologie der Salmonella- Gruppe, Kopcnhagen, 1941, 264). Habitat: Xot reported from other sources as yet. 72. Salmonella sp. (Type Berta). {Sahnonella berta Hormaeche, Peluffo and Salsamendi, Arch. Urug. de Med., Cirug. y Espec, 12, 1938, 277.) Named in honor of Prof. Arnoldo Berta, Uru- guay. Antigenic structure: IX, XII: f, g, t:— . Source: Isolated from the mesenteric glands of normal hogs. Habitat: Causes gastroenteritis in man. Also found in chickens. 73. Salmonella sp. (Type Pensacola). {Salmonella pensacola Moran and Ed- wards, Proc. Soc. Exper. Biol, and Med., 59, 1945, 52.) Antigenic structure: IX, XII: g, m, t: — . Source: From a severe case of gastro- enteritis in man. Habitat: Not reported from other sources as yet. 74. Salmonella sp. (Type Claiborne). {Salmonella claibornei Wilcox and Len- nox, Jour. Immunol., 49, 1944, 71.) Antigenic structure: I, IX, XII: k: 1,5. .. . Source: Culture isolated from human feces at Camp Claiborne, Louisiana. Habitat: Not known from other sources as yet. 75. Salmonella sp. (Type Sendai). (K type, Shimojo, quoted from Kauff- mann, Die Bakteriologie der Salmonella- Gruppe, Kopenhagen, 1941, 265; Atypi- cal Paratyphosus A, Aoki and Sakai, Cent. f. Bakt., I Abt., Grig., 95, 1925, 152; Sendai type, White, Med. Res. Council, Spec. Rept. Ser. No. 103, 1926, 118; Salmonella sendaiensis Haupt, Ergebnisse der Hyg., 13, 1932, 673; Salmonella sendai Schiitze et al., Jour. Hyg., 34, 1934, 345; Eberthella sp. (Sendai Type) F. Smith, in Manual, 5th ed., 1939, 464.) Antigenic structure: [I], IX, XII: a: 1,5.... Source : Isolated in 1922 by K. Shimojo in Japan from a case of paratyphoid. Later isolated by Aoki and Sakoi from feces, urine and blood of typhoid pa- tients. Habitat: A natural pathogen of man causing enteric fever. FAMILY EXTEROBACTERIACEAE 519 76. Salmonella sp. (Type Miami). (Salmonella niiami Edwards and Moran, Jour. Bact., 50, 1945, 259.) Antigenic structure: IX, XII: a: 1, 5 Differ culturally and biochemically from organisms of Sendai Type (Ed- wards and Moran, Jour. Bact., 50, 1945, 257). Source : Twenty-four cultures isolated by Mrs. Mildred Galton in Florida. Fourteen cultures were from cases of acute gastroenteritis, one from a patient with chronic diarrhoea, 4 from food handlers, 4 from chimpanzees thought to be affected with bacillary dysentery and one from pickles which caused an outbreak of food poisoning. One cul- ture was from Borman, Wheeler, West and Mickle (Amer. Jour. Pub. Health, 33, 1943, 127) and was isolated from a case of gastroenteritis in Connecticut. Another culture was from Seligmann, Saphra and Wassermann (Amer. Jour. Hyg., 38, 1943, 225) and was isolated from a case of enteric fever. Habitat: Api^arently widely dis- tributed as a natural pathogen of man and apes. 77. Salmonella sp. (Type Durban). (Salmonella durban Henning, Rhodes and Gordon-Johnstone, Onderstepoort Jour. Vet. Sci. An. Ind., 16, 1941, 103; also see Kauffmann, Acta Path, et Microbiol. Scand., 19, 1942, 523.) Antigenic structure: IX, XII: a: e, n, Zi5. . . . Source: Isolated by Dr. J. Gordon- Johnstone in Durban, So. Africa from feces of a woman affected with gastro- enteritis. Habitat: Not reported from other sources as yet. 78. Salmonella sp. (Type Onarimon). {Salmonella onarimon Kisida, Kitasato Arch, of Exper. Med., 17, 1940, 1.) Antigenic formula: I, IX, XII: b: 1,2 Source: From the feces of a paraty- phoid B carrier. Later found in other cases of enteric fever resembling typhoid. Habitat: Cause of a typhoid-like disease in man. 79. Salmonella sp. (Type Eastbourne). (Salmonella eastbourne Leslie and Shera, Jour. Path, and Bact., 34, 1931, 533.) Antigenic structure: [I], IX, XII: e, h:l,5 May or may not produce indole (Kauff- mann, Die Bakteriologie der Salmonella- Gruppe, 1941, 12.) Source: From human enteric fever at Eastbourne, England. Habitat: A natural pathogen for man. Also found in turkeys. 80. Salmonella sp. (Type Panama). (Jordan, Amer. Jour. Trop. Med., 14, 1934, 27; Salmonella panama Kauff- mann, Cent. f. Bakt., I Abt., Orig., 132, 1934, 160.) Antigenic structure: I, IX, XII: 1, v: 1,5.... Source: From human food poisoning at Fort Amador, in Panama, Canal Zone. Also isolated in New York City, Germany and Uruguaj-. Also in rep- tiles, hogs and chickens (Edwards and Bruner, Jour. Inf. Dis., 73, 1943, 64). Habitat: Apparently widely dis- tributed. 81. Salmonella sp. (Type Dar es Sa- laam). (Brown, Duncan and Henrj', Lancet, 1, 1926, 117; Dar-es-Salaam Typus, Schutze, Arch. f. Hyg., 100, 1928, 192; Salmonella daressalaamensis Haupt, Ergebnisse der Hyg., 13, 1932, 673; Sal- monella dar-es-salaam Schutze et al., Jour. Hyg., 34, 1934, 346.) Antigenic structure: I, IX, XII: 1, w: e, n. . . . Liquefies gelatin (Jordan, Jour. Inf. Dis., 55, 1936, 126). Source : Isolated by Butler in 1922 from a case of pyrexia at Dar es Salaam, East Africa. Cultures have also been re- ported from Zanzibar. 520 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat: Known thus far from human sources only. 82. Salmonella sp. (Type Goettingen). {Salmonella goctlingen Hohn, Cent. f. Bakt., I Abt., Orig., I46, 1940, 218.) Antigenic structure: IX, XII: 1, v: e, n, Z15. . . . The complete formula was developed by Kauffmann (Acta Path, et Microbiol. Scand., 17, 1940, 429.) Source: Not given. Presumably from a human source. Habitat: Not reported. 83. Salmonella sp. (Type .lava). (Salmonella javiana Alley and Pijoan, Yale Jour. Biol, and Med., 15, 1942, 229; Edwards and Bruner, Jour. Immunol., 44, 1942, 319.) Antigenic structure: [I], IX, XII: 1, Z28: 1,5 Source: From Eijkman Institute in Java. Isolated from feces of a child. Subsequently two cultures labeled N112 and N140, isolated in Panama from hu- man carriers, were received from Col. Chas. G. Sinclair. Habitat : Reported as yet from human sources only. 84. Salmonella gallinarum (Klein) Bergey et al. {Bacillus gallinarum Klein. Cent.f. Bakt., 5, 1889, 689; Pheasant bacillus, Klein, Jour. Path. & Bact., 3, 1893, 214; Bacillus phasiani septiciis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 3, 1896, 410; Bacterium sangui- nariiim, Moore, 12th ard 13th Ann. Rpt. for 1895-96, U. S. Dept. Agr. Bur. An. Ind., 1897, 188; see Moore, U. S. Dept. Agr. Bur. An. Ind., Bull. 8, 1895, 63; Bacillus -phasiani Migula, Syst. der Bakt., 2, 1900, 769; Bacterium phasiani septicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74; Bacterium, gallinarum Chester, ihid., 80; Bacterium pyogenes sanguinarium Berry and Ernst, Jour. Med. Res., 10 (N. S. 5), 1903-04, 402; Bacillus pseudo-cholerae gallinarum Trincas, Giorn. della R. Soc. Ital. d'Igiene, 1908, 385; Bacillus typhi galli- narum alcalifaciens and Bacillus typhi gallinarum Pfeiler and Rehse, Mitt. K. Inst. f. Landw. Bromberg, 5, 1913, 306; Eberthella sanguinaria Bergey et al., Manual, 1st ed., 1923, 231 ; Bergey et al., Manual, 2nd ed., 1925, 236; Shigella galli- narum Weldin, Iowa State Coll. Jour. Sci., 1, 1927, 179.) From Latin, of chickens. Bacterium jeffersonii Hadley, Elkins and Caldwell, Rhode Island Agr. E.xp. Sta. Bull. 174, 1918, 169 {Eberthella jeffersonii Bergey et al.. Manual, 1st ed., 1923, 230; Shigella jeffersonii Bergey et al . , Manual , 4th ed . , 1934 , 394) . Shigella jeffersonii is identical serologically with Salmonella gallinarum (St. John-Brooks and Rhodes, Jour. Path, and Bact., 26, 1923,433). Rods: 0.4 to 0.0 by 0.8 to 1.6 microns, with rounded ends, occurring singly or (in blood) in short chains. Non-motile. G ram -negative . Gelatin colonies: Small, grayish -white, finely granular, circular, entire. Gelatin stab: Slight, grayish-white surface growth with slight grayish, fili- form growth in stab. No liquefaction. Agar colonies: Moist, grayish, circular, entire. Agar slant: Thin, gray streak, with irregular margin, moist, glistening. Broth: Turbid with heavy, flocculent sediment. Litmus milk: Reaction unchanged, Ijecoming translucent. No coagulation. Potato: Slight grayish growth. Indole not formed. Nitrites produced from nitrates. Acid but no gas from glucose, fructose, galactose, mannose, xylose, arabinose, maltose, dextrin, mannitol, dulcitol and isodulcitol. Lactose, sucrose, glycerol, salicin and sorbitol are not attacked. Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide is sometimes formed. Aerobic, facultative. Optimum temperature 37°C. Antigenic structure: [I], IX, XII: — : — . Identical with Salmonella pullorum,, FAMILY r:NTEROBACTERIACEAE 521 and related to Salmonella iyphosa. [I] antigen noted by Kauffmann (Acta Path, et Microbiol. Scand., Suppl. 54, 1944, 36). Source and habitat: The causative agent of fowl typhoid (clearly to be dis- tinguished from fowl cholera), and identical with Moore's infectious leu- kemia of fowls. Infectious for rabbits and all poultry, canaries and certain wild birds (quail, grouse, pheasant) by feeding or by injection. Found once in a normal human carrier. 85. Salmonella pullorum (Rettger) Bergey et al. (Bacterium pullorum Rettger, Jour. Med. Res., ^7 (N.S. 16), 1909, 117; also see Rettger, X. Y. Med. Jour., 71, 1900, 803; ibid., 73, 1901, 267; Rettger and Harvey, Jour. Med. Res., 18 (N.S. 13), 1908, 277; Bacillus pullorum. Smith and Ten Broeck, Jour. Med. Res., 31 (N. S. 26), 1915, 547; Bergey et al.. Manual, 1st ed., 1923, 218; Typus pul- lorum, Kauffmann, Zentbl. f. d. ges. Hyg., £5, 1931, 273.) From Latin, of chickens. Rods: 0.3 to 0.5 by 1.0 to 2.5 microns, occurring singly. Xon-motile. Gram- negative. Gelatin colonies: Grayish-white, moist, lobate, with grape-leaf surface. Gelatin stab: Slight, grayish surface growth. No liquefaction. Agar colonies: Grayish-white, smooth, glistening, entire to undulate. Agar slant: Develops as discrete, translucent colonies. Broth: Turbid. Litmus milk: Acid, becoming alkaline. No coagulation. Potato: Slow development, grayish. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, arabinose, xylose, mannitol and rhamnose. Does not at- tack lactose, sucrose, maltose, dextrin, salicin, raffinose, sorbitol, adonitol, dulcitol or inositol. Gas may be slight or absent (cf. Salmonella gallinarum). Xylose may be fermented late (see Wel- din, Iowa State Coll. Jour. Sci., 1, 1927, 165K ^Maltose fermenting strains may occur (Hinshaw, Brown© and Taylor, Jour. Inf. Dis., 73, 1943, 197). Reduces trimethylamine oxide (Wood and Baird, loc. cit.). Hydrogen sulfide is formed. Aerobic, facultative. Optimum temperature 37°C. Antigenic structure, IX, XII: — : The complete antigenic formula of S. pullorum is IX, XIL, XIL, XII3, while that of S. gallinarum seems to be IX, XIIi, XII3. Antigen XIIo is variable in S. pullorum (Edwards and Bruner, Cor- nell Vet., 36, 1946, 318) and XII2++ and XII2+ forms occur. The XII2+- forms are synonj^mous with the X strains of Younie (Can. Jour. Comp. Med., 5, 1941, 164). Source: Isolated from chickens and other birds, as well as calves, hogs, rab- bits and man. Occasionally produces food poisoning or gastroenteritis in man (Mitchell, Garlock and Broh- Kahn, Jour. Inf. Dis., 79, 1946, 57). Habitat: The cause of white diarrhoea in young chicks. Infects the ovaries and eggs of adult birds. 85a. Salmonella f/allinarum var. Duis- burg. (Miiller, IMtinch. med. Wchnschr., 80, 1933, 1771 ; Kauffmann, Cent. f. Bakt., I Abt., Orig., 132, 1934, 337.) Antigenically identical with Salmo- nella gallinarum ixiid Salmonella pullorum. Differs from Salmonella gallinarum in its slow fermentation of maltose, failure to ferment d-tartrate and in not forming H2S. Source and habitat: Isolated from acute gastroenteritis in man. 86. Salmonella sp. (Type Canastel). {Salmonella canastel Pcandall and Bruner, Jour. Bact., Jf9, 1945, 511.) Source of name not given. Liquefies gelatin. Antigenic structure: IX, XII: Z29: 1,5.... 522 MANUAL OF DETERMINATIVE BACTERIOLOGY Source: Isolated in North Africa from American soldiers acting as food hand- lers. Habitat: Not reported from other sources as yet. 87. Salmonella sp. (Type Italia). {Salmonella italiana Bruner and Ed- wards, Proc. Soc. Exp. Biol, and Med., 58, 1945, 289.) Antigenic structure: IX, XII: 1, v: 1, 11 Source: Two cultures, one isolated from a case of bloody diarrhoea and the other from a case of gastroenteritis in man. Found in Italy by Lt. Col. Robert Hebble and by Capt. Ira C. Evans. Habitat: Not reported from other sources as j^et. S8. Salmonella sp. (Tj^pe Napoli). {Salmonella napoli Bruner and Edwards, Proc. Soc. Exper. Biol, and Med., 58, 1945,289.) Antigenic structure: [I], IX, XII: 1, Z13: e, n, X. . . . Source: Ten cultures isolated from normal feces and from cases of gastro- enteritis in Naples, Italy. The first culture was isolated by Capt. W. H. Ewing. Habitat: Not reported from other sources as j'et. 89. Salmonella sp. (Tj'pe Loma Linda) . {Salmonella, loma linda Edwards, Proc. Soc. Exper. Biol, and Med., 57, 1944, 104.) Antigenic structure: IX, XII: a: e, n, X. . . . Source: Single culture isolated by Dr. T. F. Judefind, Loma Linda, California from the spinal iluid of a baby that died of meningitis. Habitat: Not reported from other sources as yet. 90. Salmonella sp. (Type New York). {Salmonella new york Kauffmann, Acta Path, et Microbiol. Scand., Suppl. 54, 1944, 35.) Antigenic structure: IX, XII: 1, v: 1,5 Source: Found by Dr. F. Schiff, New York in a study of a culture received under the label S. panama Strain No. 431. Regarded at the present time as a strain of Salmonella javiana by Dr. Kauffman (personal communication, March, 1947). Habitat: Not reported from other sources as yet. 91. Salmonella sp. (Tj^pe London). {Salmonella Type L, White, Med. Res. Council Spec. Rept. Ser. 103, 1926, 37;Sal- monella londonensis Haupt, Ergebnisse der Hyg., 13, 1932, Q7 5; Salmonella london Schtitze et al., Jour. Hyg., 34, 1934, 346.) Antigenic structure: III, X, XXVI: I, v: 1,6 Source: Lsolated in Loudon from the feces of a gastroenteritis patient ^'rom Reading, England. Habitat: Found in human infections, in hogs and in chickens. 92. Salmonella sp. (Type Give) . {Sal- monella give Kauffmann, Ztschr. f. Hyg., 120, 1937, 177.) Antigenic structure: III, X, XXVI: 1, v;l, 7 Source: From feces of a patient with pernicious anemia. Also found in the U. S. A. and Germany. Occurs in fowls and hogs (Edwards and Bruner, Jour. Inf. Dis., 72, 1943,64). Habitat: Apparently widely dis- tributed. 93. Salmonella sp. (Type Uganda). {Salmonella Uganda Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940, 189.) Antigenic structure: III, X, XXVI: 1, Z13: 1, 5. . . . Source: Isolated in Uganda by Dr. H. G. Wiltshire from a human spleen on autopsy. Tyjied by Dr. F. Kauff- mann. Habitat : Not reported from other sources as yet. FAMILY ENTEROBACTERIACEAE 523 94. Salmonella anatis (Rettger and Scoville) Bergej' et al. {Bacterium anatis Rettger aud Scoville, Abst. Bact., 3, 1910, S; not Bacterium anatis Migula, Syst. d. Bakt., 2, 1900, 364; Bacteriuvi anatum Rettger and Scoville, Jour. Inf. Dis., 26, 1920, 217; Escherichia anata Bergey et al., Manual, 1st ed., 1923, 198; Bergey et al., IManual, 2nd ed., 1925, 238; Salmonella anatum Bergey et al., Manual, 3rd ed., 1930, 344.) From Latin, of the duck. With the transfer of this organism to the genus Salmonella, the original species name anatis again becomes available in spite of the earlier use of this species name by ]\Iigula for Cornil and Toupet's Bacillus der Enten-cholera (Compt. rend. Acad. Sci., Paris, 106, 1888, 1737). The latter organism is stated bj' Rettger and Sco\alle (1920, loc. cii., 220) to be indistinguishable from Pasteurella avi- septica. Morphologj^ and cultural characters like those of Salmonella enteritidis. Kauffmann (Ztschr. f. Hyg., 119, 1937, 352) describes a lactose-splitting variant of this species. Antigenic structure: III, X, XXVI: e, h: 1, 6. . . . Reduces trimethylamine oxide (Wood and Baird {loc. cit.). Source: Isolated from an epizootic of keel in ducklings. Also found in in- testinal infections in chickens and man. Frequently occurs in association with Salmonella typhimiiriiim. Habitat : Widely distributed in man and domestic animals. 95. Salmonella sp. (Tj^pe Muenster). {Salmonella anatum var. muenster, Kauff- mann and Silberstein, Cent. f. Bakt., I Abt., Orig., 132, 1934, 431; Salmonella m,uenster Kauffmann, Ztschr. f. Hyg., HO, 1937, 177.) Antigenic structure: III, X, XXVI: e, h: 1, 5. . . . Source: Isolated by Dr. Besserer in Muenster from food poisoning. Also isolated in Uruguay from human sources. Habitat: Xot known from any but human sources as yet. 96. Salmonella sp. (Type Xyborg). {Salmonella anatum, var. nyborg, Kris- tensen and Bojlen, Cent. f. Bakt., I Abt., Orig., 136, 1936, 294; Salmonella nyborg Kauffmann, Ztschr. f. Hyg., 120, 1937, 189.) Antigenic structure: III, X, XXVI: e, h: 1, 7. . . . Source : From a case of acute enteritis in a young girl in Nyborg, Denmark. Habitat: Ivnown only from human sources as yet. 97. Salmonella sp. (Type Vejle). {Sal- monella vejle Harhoff , quoted from Kauff- mann, Die Bakteriologie der Salmonella- Gruppe, Kopenhagen, 1941, 274.) Antigenic structure: III, X, XXVI: e, h:l,2, 3. . . . Source: Isolated by E. M0ller, Copen- hagen, from a case of acute gastro- enteritis. Habitat: Xot reported from other sources as yet. 98. Salmonella sp. (Type Meleagris). {Salmonella meleagridis Bruner and Edwards, Amer. Jour. Hyg., 3J^, 1941, 82; not Salmonella meleagridis Rettger, Plastridge and Cameron, Jour. Inf. Dis., 53, 1933, 279.) Antigenic structure: III, X, XXVI: e, h: 1, w. . . . Source: Original cultures isolated by Dr. B. S. Pomeroy, Univ. of Minnesota, from two distinct outbreaks of infection in turkey poults. Stated to be the same as Salmonella bantam from Batavia, Java (Kauffmann, Acta Path, et Mi- crobiol. Scand., 19, 1942, 529). Habitat : In addition to the two strains isolated in Minnesota (Bruner and Ed- wards, Kentucky Agr. Exp. Sta., Bull. 434, 1942), the same type was recognized among cultures received from Massa- chusetts, Michigan, Pennsylvania, ^Maryland, South America and Japan. Also isolated from German soldiers in 524 MAXUAi. OF dp:termixative bacteriology Norway by Tesdal (Kauffmann, Die Bakteriologie der Salmonella-Gruppe, 1941, 295) and from snakes by Hinshaw and McNeil (Amer. Jour. Vet. Res., 6, 1945,264). 99. Salmonella sp. (Type Shangani). {Salmonella shangani Kauffmann, Acta Path, et Microbiol. Scand., 16, 1939, 347.) Antigenic structure: III, X, XXVI: d: 1,5.... Source: Isolated in Zanzibar by Dr. J. D. Robertson from a woman with enteric fever. Habitat: Known only from human sources as yet. 100. Salmonella sp. (Type Zanzibar). (Salmonella Zanzibar Kauffmann, Acta Path, et Microbiol. Scand., M, 1939,347.) Antigenic structure: III, X, XXVI: k: 1, 5. . . . Source: Isolated in Zanzibar by Dr. J. D. Robertson from a typhoid carrier. Habitat: Also found in chickens (Edwards) . 101. Salmonella sp. (Type Amager). {Salmonella amager Kauffmann, Acta Path, et Microbiol. Scand., 16, 1939, 347.) Antigenic structure: III, X, XXVI: y: 1, 2, 3. . . . Source: Isolated in Copenhagen from the feces of a person suffering from gas- troenteritis. Habitat: Known only from human sources as yet. 102. Salmonella sp, (Type Lexington). {Salmonella lexington Rubin, Jour. Bact., 40, 1940, 463; Edwards, Bruner and Rubin, Proc. Soc. Exper. Biol, and Med., U, 1940, 395.) Antigenic structure: III. X, XXVI: zio: 1,5.... According lo Kauffmann (Die Bak- teriologie der Salmonella-Gruppe, 1941, 276), Dr. Erber of Java has found a Salmonella type with the same antigenic structure and has given it the name Salmonella batavia. Source: Isolated from mesenteric lymph glands of apparently normal hogs by Dr. H. L. Rubin, Univ. of Kentucky, Ijcxington, Ky. Habitat: Also reported from turkeys. 103. Salmonella sp. (Type Weltevre- den). {Salmonella iveltevreden Mertens, quoted from Kauffmann, Acta Path, et Microbiol. Scand., 19, 1942, 529.) Antigenic structure: III, X, XXVI: r: Z6. . . . Source : Isolated by Dr. W. K. Mertens, Batavia, Java, according to Kauffmann {loc. cit.). Habitat: Not recorded in available literature. 104. Salmonella sp. (Type Orion). {Salmonella type, var. orion and Sal- monella orion Barnes, Cherry and Myers, Jour. Bact. 50, 1945, 578.) From a sea- man on the S. S. Orion. Antigenic structure: III, X, XXVI: y:l, 5. ... Source: From rectal swab specimen from a normal food handler. Habitat: Not reported from other sources as yet. 105. Salmonella sp. (Type Butantan). {Salmonella butantan Peluffo, Arch. Urug. de Med., Cirug. y Espec, 18, 1944,000.) Antigenic structure : III, X, XXVI : b : 1,5 Source: Isolated by Dr. C. A. Peluffo from a case of diarrhoea in a child. Habitat: Not reported from other sources as yet. 106. Salmonella sp. (Type Newington). (Anatum Ci No. 3071, N.C.T.C, London, Kauffmann and Silberstein, Cent. f. Bakt., I Abt., Orig., 132, 1934, 434; Salmonella newington Edwards, Jour. Hyg., 37, 1937, 384.) Antigenic structure: III, XV: e, h: 1,6 Source: Isolated from ducks from Newington, Connecticut by Dr. L. F. FAMILY ENTEROBACTERIACEAE 525 Rettger. Also found in hogs, silver foxes and man. Kauffmann (Ztschr. f. Hyg., 120, 1937. 177) has described a re- lated type (Salmonella lim) from a case of enteritis in Tim, Denmark. Habitat : Widely distributed. 107. Salmonella sp. (Type Selandia). (Salmonella selandia Kauffmann, Ztschr. f. Hyg., 7:20, 1937, 189.) Antigenic structure: III, XV: e, h: 1, 7. . . . Source: Isolated from the feces of a sailor on the S. S. Selandia after a voy- age to Asia and Australia. Was patient in Bispebjerg Hospital with pleuro- pneumonia at the time. Habitat: Known only from human sources as yet. 108. Salmonella sp. (Type New Bruns- wick). (Salmonella new bnmsivick Ed- wards, Jour. Hyg., 37, 1937, 384; also see Kauffmann, Ztschr. f. Hyg., 120, 1937, 189.) Antigenic structure: III, XV: 1, v: 1, 7. . . . Source: Isolated by Dr. F. R. Beau- dette. New Brunswick, New Jersey from a chicken. Also isolated from gastro- enteritis in man. Habitat: Apparently widely dis- tributed. 109. Salmonella sp. (Type Illinois). (Salmonella illinois Edwards and Bruner, Proc. Soc. Exper. Biol, and Med., 48, 1941, 240.) Antigenic structure: (III), (XV), XXXIV: zio: 1, 5 Source: Isolated from hogs in Illinois by Dr. Robert Graham, from Hungarian partridges in Michigan by Miss Virginia Stoney and from turkeys in Minnesota by Dr. B. S. Pomeroy. Habitat: Also reported from hogs and man (Edwards). 110. Salmonella sp. (Type Senften- berg). (Typus Senftenberg, Kauff- mann, Ztschr. f. Hyg., Ill, 1930, 221; Salmonella senftenberg Schiitze et al., Jour. Hyg., 54, 1934, 339; Salmonella senftenbergensis Haupt, Ergebnisse der Hyg., 13, 1932,673.) Antigenic structure: I, III, XIX: g, s, t: — . Source: From a case of acute gastro- enteritis in a boy in Senftenberg, Den- mark. Cultures have frequently been found from persons and also from young turkeys. Habitat: Apparently widely dis- tributed. 111. Salmonella sp. (Type Niloese). (Salmonella niloese Kauffmann, Acta path, et Microbiol. Scand., 16, 1939, 347.) Antigenic structure: I, III, XIX: d: Z6. . . . Source: Isolated in Copenhagen from a case of acute gastroenteritis in Niloese, Denmark. Later found frequently in gastroenteritis in Denmark. Habitat: Known only from human sources as yet. 112. Salmonella sp. (Type Simsbury). (Salmonella simsbury Bruner and Ed- wards, Proc. Soc. Exper. Biol, and Med., 50, 1942, 174.) Antigenic structure: I, III, XIX: Z27: Source: Original culture isolated by Dr. E. K. Borman, State Dept. Health Lab., Hartford, Conn., from a normal human carrier from Simsbury, Conn. Edwards states (1946) that this may be a variant of Salmonella sp. (Type Senf- tenberg) . Habitat: Also found in turkeys (Bru- ner and Edwards, Kentucky Agr. Exp. Sta., Bull. 434, 1942. 9). 113. Salmonella sp. (Type Taksony). (Sabnonclla taksony Rauss, Ztschr. f. Immunitatsforsch., 103, 1943, 220.) Antigenic structure: I, III, XIX: i: Z6. . . . Source : Isolated from a healthy carrier (Hungary). 526 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat: Not repoi'ted from other sources as yet. 114. Salmonella sp. (Type Kentucky). (Salmonella kenlucky Edwards, Jour. Hyg., 38, 1938, 306.) Antigenic structure: (VIII), XX: i: Z6. . . . Source: Isolated from the intestinal tract of a chick affected with coccidiosis and ulcerative enteritis. Found at Le.xington, Kentucky. Habitat: Also reported from many species of fowls, from hogs and from man (Edwards). 115. Salmonella sp. (Type Aberdeen). (Salmonella aberdeen J. Smith, Jour. Hyg., 34, 1934, 357.) Antigenic structure: XI: i: 1, 2, 3. . . . Source: Isolated in Aberdeen, Scot- land, from the stool of a child suffering from acute enteritis. Also isolated by Timmerman in Utrecht from Ovomal- tine, and by Edwards in Kentucky from birds. See Kauffmann, Die Bakteriol- ogie der Salmonella-Gruppe, Kopen- hagen, 1941, 279. Habitat: Apparentlj^ widely dis- tributed. 116. Salmonella sp. (Type Rubislaw). (Salmonella rubislaw Smith and Kauff- mann, Jour. Hyg., ^0, 1940, 122.) Antigenic structure: IX: r: e, n, x. . . . Source: Isolated in Aberdeen, Scot- land from the feces of a child suffering from enteritis. Also found by Tesdal in Oslo, Norway. Reported by Hin- shaw and McNeil from snakes (Amer. Jour. Vet. Res., 6, 1945, 264). Habitat: Apparenth' widely dis- tributed. 117. Salmonella sp. (Type Pretoria). (Salmonella pretoria Henning, Rhodes and Gordon -Johnstone, Onderstepoort Jour. Vet. Sci. An. Ind., 16, 1941, 103.) Antigenic structure: XI: k: 1,2, 3. . . . Source : Isolated by Dr. M. W. Henning in Pretoria, South Africa from an infec- tion in garbage-fed hogs. Habitat: Not reported from other sources as yet. 118. Salmonella sp. (Type Venezia). (Salmonella veneziana Bruner and Joyce, Jour. Bact., 50, 1945, 371.) Antigenic structure: XI: i: e, n, x. . . . Source: Culture received from Capt. J. K. Hill. Isolated from an apparently normal Italian civilian food handler in Venice, Italy. Habitat: Not known from other sources as yet. 119. Salmonella sp. (Type Solt)r (Sal- monella soli Rauss, Ztschr. f. Immuni- tatsforsch., 103, 1943, 220.) Antigenic structure: XI: y: 1, 5. . . . Source : Isolated from a healthy carrier (Hungary) . Habitat: Not reported from other sources as yet. 120. Salmonella sp. (Type St. Lucie). (Salmonella luciana Moran, Edwards and Bruner, Proc. Soc. Exp. Biol, and Med., 64, 1947, 89.) From St. Lucie, Florida. Antigenic structure: XI: a: e, n, Zis. . . . Source : Single culture isolated by Mrs. Mildred Galton from feces of a normal human carrier. Habitat: Not known from other sources as yet. 121. Salmonella sp. (Type Senegal). (Salmonella Senegal Hinshaw and McNeil, Jour. Bact., 52, 1946, 349.) Antigenic structure: XI: r: 1, 5. . . . Source: Isolated by Dr. W. L. Hin- shaw from a green mamba snake. Habitat: Not known from other sources as yet. 122. Salmonella sp. (Type Marseille). (Salmonella marseille Moran, Edwards and Bruner, Proc. Soc. Exp. Biol, and Med., 64, 1947, 89.) FAMILY ENTEROBACTERIACEAE 527 Antigenic structure: XI: a: 1, 5. . . . Source: Isolated in Marseilles, France by Capt. Wm. Sutton from feces. Habitat: Not known from other sources as yet. 123. Salmonella sp. (Type Grumpy). (Salmonella grumpensis Hormaeche and Peluffo, quoted from Hormaeche et al., Jour. Bact., 47, 1944, 323.) Xamed for a person called grumpj'. Antigenic structure: XIII, XXIII, XXXVI: d: 1, 7 ... as given by Kauff- mann (Acta Path, et ^Microbiol. Scand., Suppl. 54, 1944, 37). Source: Isolated in Uruguay from a guinea pig. Also studied by Kauffmann {Joe. cit.). Habitat: Xot reported from other sources as j'et. 124. Salmonella sp. (Type Poona). (Salmonella poona Bridges and Scott, Jour. Roy. Army Med. Corps, 55, 1935, 221.) Antigenic structure: XIII, XXII: z: 1, 6. . . . Source: Isolated by Dr. L. Dunbar in Poona from the stool of a child suffering from enteritis. Habitat: Also reported from hogs (Edwards) . 125. Salmonella sp. (Type Borbeck). (Salmonella borbeck Hohn and Herri - mann, Cent. f. Bakt., I Abt., Orig., 145, 1940, 219.) Antigenic structure: XIII, XXII: 1, v: 1, 6. . . . Source: Isolated from the feces of a child with typhoid. Found in the Bor- beck section of Essen, Germany. Habitat: Xot reported from other sources as yet. Source: Isolated by the State Dept. of Health of Mississippi from the stool of a normal food handler. Habitat: Also reported from hogs (Edwards). 127. Salmonella sp. (Type Wichita). (Salmonella wichita Schiff and Strauss, Jour. Inf. Dis., 65, 1939, 125.) Antigenic structure: I, XIII, XXIII: d:— . Source: Isolated by ^liss B. McKinlay in an epidemic of enteritis affecting babies, \Yichita, Kansas. Also in fowls, turkej's and hogs (Edwards and Bruner, Jour. Inf. Dis., 72, 1942, 64). Habitat: Apparently widely dis- tributed. 128. Salmonella sp. (Type Havana). (Salmonella havana Schiff and Saphra, Jour. Inf. Dis., 68, 1941, 125.) Antigenic structure: I, XIII, XXIII: Source: Isolated during an outbreak of 21 cases of meningitis in children in a maternitj^ hospital in Havana, Cuba. Habitat: Xot reported from other sources as yet. 129. Salmonella sp. (Type Worthing- ton). (Salmonella worthington Edwards and Bruner, Jour. Hyg., 38, 1938, 716.) Antigenic structure: I, XIII, XXIII: 1, w: z. . . . Source: Isolated by Dr. B. S. Pomeroy from a turkey poult from Worthington, Minnesota. Also found in a hen. Later additional cultures were found in other birds, in rodents, cattle, hogs and man. (Edwards and Bruner, Jour. Inf. Dis., 72, 1943,64). Habitat: Apparently widely dis- tributed. 126. Salmonella sp. (Type Mississippi) . (Salmonella mississippi Edwards, Cherry and Bruner, Proc. Soc. Exp. Biol, and :Med., 54, 1943, 263.) Antigenic structure: I, XIII, XXIII, b: 1, 5. . . . 130. Salmonella sp. (Type Cuba). (Salmonella cubana Seligmarm, Wasser- man and Saphra, Jour. Bact., 51, 1946, 123.) Antigenic structure: I, XIII, XXIII: Z29: — ■ 528 MANUAL OF DETERMTXATIVE BA('TEKIOT>OGY iSource: Isolated in Havana,, Cuba by Dr. Arturo Curbelo from diseased baby chicks. Habitat: Not reported from other sources as yet. 131. Salmonella sp. (Type Heves). (Salmonella heves Rau.ss, Ztschr. f. Im- munitatsforsch., 103, 1943, 220.) Antigenic structure: VI, XIV, XXIV: d: 1, 5 Source: Isolated from a healthy carrier (Hungary) . Habitat: Not reported from other sources as yet. 132. Salmonella sp. (Type Carrau). (Salmonella carrau Hormaeche, Peluffo and Salsamendi, Arch. Urug. de Med., Cirug. y Espec, 12, 1938, 377; Hor- maeche, Peluffo and Pereyra, Jour. Bact., 47, 1944, 323.) Antigenic structure: VI, XIV, XXIV: y:l, 7. ... Source: Isolated in Uruguay from mesenteric glands of normal hogs. Habitat: Also reported from feces and blood in man, once from flies and one culture from human blood from Mexico. Source: Isolated by Mrs. Mildred Gal- ton from feces of a patient with a febrile disease and diarrhoea. Habitat: Also reported from reptiles (Edwards). 135. Salmonella sp. (Type Madelia). (Sahnonella madelia Cherry, Edwards and Bruner, Proc. Soc. Exp. Biol, and Med., 52, 1943, 125.) Antigenic structure: (I), VI, XIV, XXV: y: 1.7.... Source: A single culture isolated by Dr. B. S. Pomeroy from the liver of a poult that died of septicemia. Found in Madelia, Minnesota. Habitat: Also reported from man (Edwards). 136. Salmonella sp. (Type Sundsvall). (Sahnonella sundsvall Olin and Alin, Acta Path, et Microbiol. Scand., 20, 1943, 607.) Antigenic structure: (I), VI, XIV, XXV: z: e, n, X. . . . Source: Isolated from a person suffer- ing from gastroenteritis. Habitat: Not reported from other sources as vet. 133. Sahnonella sp. (Type Onderste- poort). (Salmonella onderstepoort Hen- ning. Jour. Hyg., 36, 1936, 525.) Antigenic structure: (I), VI, XIV, XXV: e, (h): 1, 5. . . . Source: Isolated in So. Africa bj' Dr. J. H. Mason from sheep in Onderste- poort. Also isolated from man by Dr. Hormaeche (Uruguay) and from turkeys (Edwards, Kentucky). Habitat: Apparently widely dis- tributed in warm-blooded animals. 134. Salmonella sp. (Type Florida). (Sahnonella florida Cherry, Edwards and Bruner, Proc. Soc. Exp. Biol, and Med., 52, 1943, 125; Galton and Quan, Amer. Jour. Hyg., 38, 1943, 173.) Antigenic structure: (I), VI, XIV, XXV: d: 1, 7. . . . 137. Salmonella sp. (Type Orient). (Salmonella orientalis Carlquist and Conte, Bull. U. S. Army Med. Dept., 6, 1946, 343.) Antigenic .structure: XVI: k: e, n, Zlo. . . . Source: Isolated from U. S. Army personnel who had been prisoners of the Japanese Army in the Orient. Habitat: Not known from other sources as yet. 138. Salmonella sp. (Type Hvitting- foss). (Salmonella hvitiingfoss Tesdal, Ztschr. f. Hyg., 118, 1936, 533.) Antigenic structure: XVI: b: e, n, Source: Isolated during a food poison- ing outbreak in Hvittingfoss, a small town in Norway. Caused by eating FAMILY ENTEROBACTERIACEAE 529 pultoste, a kind of soft cheese. Cul- tures secured from the cheese, from the persons who were poisoned, ffom sewage and from a foal. Habitat: Evidently rather widelj' distributed. 139. Salmonella sp. (Type Gaminara). (Salmonella gaminara Hormaeche, Pe- luffo and Salsamendi, Arch. Urug. de Med., Cirug. y Espec, 12, 1938, 377; ibid., 14, 1939, 217.) Named in honor of Prof. Gaminara of Uruguay. Antigenic structure: XVI: d: 1, 7. . . . Source: Isolated from the feces of a child suffering from enteritis. Habitat: Xot known from other sources as yet. 140. Salmonella sp. (Type Szentes). {Salmonella szentes Rauss, Ztschr. f. Immunitatsforsch., 103, 1943, 220.) Antigenic structure: XVI: k: 1, 2, 3 Source: Isolated by Dr. K. Rauss from a healthy carrier (Hungary) . Habitat: Not reported from other sources as yet. 141. Salmonella sp. (Type Kirkee). (Salmonella kirkee Bridges and Dunbar, Jour. Roy. Army Med. Corps, 67, 1936, 289.) Antigenic structure: XVII: b: 1,2. . . . Source: Isolated in Kirkee, India from the feces of a child suffering from acute enteritis. The source of the iufection was thought to be a dog. Habitat: Not reported from other sources as yet. 142. Salmonella sp. (Type Cerro). (Bacterium cerro Hormaeche, Peluffo and Salsamendi, Arch. Urug. ^led., Cirug. y Espec, 12, 1938, 377; Salmonella cerro Hormaeche, Peluffo and Aleppo, ihid., 19, 1941, 125.) Antigenic structure: XVIII: zn, z^s, Zoil — . Source: Isolated from the mesenteric glands of normal hogs from Cerro, Uruguay. Habitat: Also isolated by the authors in 13 cases of infantile infections. Found also in chickens (Edwards). 143. Salmonella sp. (Type Minnesota) . (Salmonella minnesota Edwards and Bru- ner. Jour. Hyg., 38, 1938, 716.) Antigenic structure: XXI, XXVI: b: e, n, X. . . . Source: Isolated in Minnesota by Dr. B. S. Pomeroy from a young turkey. Habitat: Also reported from rattle and man. 144. Salmonella sp. (Type Tel Aviv). (Salmonella tel-aviv Kauffmann, Acta Path, et Microbiol. Scand., 17, 1940, 1.) Antigenic structure: XXVIII: y: e, n, Zi5. . . . Source: Isolated in Tel Aviv, Palestine by Dr. G. B. Simmins during an epi- zootic affecting young chickens during which 50 per cent died. Habitat: Not known from other sources as yet. 145. Salmonella sp. (Type Pomona). (Salmonella pomona Edwards, Proc. Soc. Exp. Biol, and Med., 58, 1945, 291.) Antigenic structure: XXVIII: y: 1, 7. . . . Source: Single culture isolated from the intestine of a poult in 1941 by Dr. W. R. Hinshaw. Habitat: Also reported from man (Edwards). 146. Salmonella sp. (Type Ballerup). (Salmonella ballerup Kauffmann and M0ller, Jour. Hyg., 40, 1940, 246.) Antigenic structure: XXIX, [Vi]: Zi4: — . Source: From the feces of a woman from the town of Ballerup, Denmark. A cause of gastroenteritis. Habitat: Not known from other sources as yet. 147. Salmonella sp. (Type Hor- maeche). (Salmonella hormaechei Mon- teverde. Nature, 154, 1944, 676.) Named in honor of Dr. Hormaeche of Uruguay. 530 MANUAL OF DKTERMIXATIVE BACTERIOLOGY Antigenic structuro: XXIX, [Vi]*: 2,30, [zsi]: — . Source : From the ovary of a hen whose blood gave a positive reaction with the »S'. pallor um antigen. Found in Buenos Aires by Dr. Monteverde. Habitat: Also reported from hogs and man (Edwards). ♦Reported by Dr. P. R. Edwards (personal communication). 148. Sahnonella sp. (Tj'pe Urbana). (SabnoticUa urbana Fldwardsand Bruner, Jour. Inf. Dis., 69, 1941, 223.) Antigenic structure: XXX: b: e, n, Source: One culture was received from Dr. Robert Graham, Urbana, Illinois and was isolated from the contents of the colon of a hog aft'ected with hemorrhagic enteritis. The second culture was iso- lated from the intestinal tract of a chicken by Dr. W. L. Mallmann, East Lansing, Michigan. Habitat: Also reported from man (Edwards) . 149. Salmonella sp. (Type Adelaide). {Salmonella adelaide Cleland. Med. Jour. Australia, 31, 1944, 59.) Antigenic structure: XXXV: f, g: — . Source : Isolated in Adelaide, Australia by Miss Nancy Atkinson from two fatal cases resembling typhoid fever. Habitat: Not reported from other sources as yet. 150. Salmonella sp. (Type Inverness). (Salmonella Inverness Edwards and Hughes, Proc. Soc. Exp. Biol, and Med., 56, 1944, 33.) Antigenic structure: XXXVIII: k: 1, 6 Source: Isolated by Mrs. Mildred Gal- ton and Mr. M. S. Quan of the Florida State Department of H*ealth, from the stool of a normal food handler, Inver- ness, Florida. Habitat: Not reported from other sources as yet. 151. Salmonella sp. (Type Cham- paign). (Salmonella champaign Ed- wards, Proc. Soc. Exp. Biol, and Med., 58, 1945, 291.) Antigenic structure: XXXIX: k: 1,5. .. . Source: Single culture isolated from the liver of an adult hen by Dr. Robert Graham, Champaign, Illinois. Habitat: Not reported from other sources as yet. Appendix I: The following species and varieties are largely taken from Haudu- roy, Ehringer, Urbain, Guillot and Magrou, Dictionnaire des Bact^ries Pathogenes, Paris, 1937, 446-472. The relationships of many of these are not clear. Bacillus canariensis Migula. (Ba- cillus der Kanarienvogelseptikamie, Rieck, Deutsche Ztschr. f. Thiermed., 15, 1889, 69; Migula, Syst. d. Bakt., S, 1900, 770; Bacillus avisepticus Chester, Man. Determ. Bact., 1901, 220; not Ba- cillus avisepticus Kitt, in Ivolle and Wassermann,Handb. d. path.Mikroorg., 1 Aufl., 2, 1903, 544.) Associated with intestinal catarrh and liver changes in canaries. Hadley, Elkins and Caldwell (Rhode Island Agr. Exp. Sta., Bull. 174, 1918, 178) regard this as probably Ba- cillus gallinarum Klein. Bacillus friedehergensis Kruse. (Ba- cillus der Friedeberger Fleischvergif- tung, Gaffky and Paak, Mitt. a. d. kaiserl. Gesundheitsamte, 6, 1890, 159; Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 378; Bacterium friede- hergensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 73.) From sausage in meat poisoning. Salmonella abortus canis Gard. (Ztschr. f. Hyg., 121, 1938, 139.) From the feces of four persons with paratyphoid appar- ently spread from an infected dog. Kauffmann regards this as identical with Salmonella schottmuelleri. . Salmonella annamensis Hauduroy et al. (Un bacille du groupe des Salmonella, Normet, Urbain and Chaillot, Compt. rend. Soc. Biol., Paris, 101, 1929, 752; Hauduroy et al., Diet. d. Bact. Path., FAMILY EN'TEROBACTERIACEAE 531 1937, 450.) Isolated during an epidemic of dysentery at Hue (Annam) in 1925. Salmonella archibaldii Castellani and Chalmers. (Man. Trop. INIed., 3rd ed., 1919, 940.) Salmonella Carolina (Castellani) Cas- tellani and Chalmers. {Bacillus caro- li?ius Castellani, Ann. di Med. Xav. e Colon., 1, 1918; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 940.) Salmonella coagulans (Castellani) Hau- duroy et al. {Bacillus coagulans Cas- tellani, 1916; Balkanella coagulans Cas- tellani, 1916; see Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 935; Hauduroy et al., Diet. d. Bact. Path., 1937, 453.) Salmonella columhensis (Castellani) Castellani and Chalmers. {Bacterium columbense Castellani, Proc. Meeting Ceylon Branch British Assoc, 1905, quoted from Castellani, Cent. f. Bakt., I Abt., Orig., 74, 1914, 197; Bacillus columhensis Castellani, Jour. Trop. Med. and Hyg., 20, 1917, 181; Castellani and Chalmers, Ann. Inst. Past., 34, 1920, 609; Morganclla columhensis Fulton, Jour. Bact., 46, 1943,81.) The cause of colum- hensis fever. Isolated from feces, urine and blood. Salmonella enteritidis var. v, Haudu- roy et al. (Bacille para-Gartner V, Rochaix and Couture, Revue de Micro- biologie appliquee, 2, 1936; Hauduroj' et al.. Diet. d. Bact. Path., 1937, 454.) Found associated with Salmonella en- teritidis in meat pies and in the feces of individuals with food poisoning. Salmonella enteritidis -yellow , a variety of Salmonella enteritidis Deskowitz and Buchbinder (Jour. Bact., 29, 1935, 293). Cultures differ from typical Salmonella enteritidis in producing a yellow, water- soluble pigment. From the feces of a rat with enteric infection. Salmonella foetida Bergey et al. {Coc- cohacillis foetidus ozenae Perez, Ann. Inst. Past., 13, 1899, 937; Coccobacillus {foetidus) ozaenae Ward, Jour. Bact., 2, 1917, 619; Bergey et al.. Manual, 1st ed., 1923, 220; Bacterium foetida Weldin and Levine, Abst. Bact., 7, 1923, 13; Es- cherichia foetida Bergey et al.. Manual, 2nd ed., 1925, 222.) From chronic rhi- nitis, ozena. See INIanual, 4th ed., 1934, 380 for a description of this species. Salmonella holsatiensis Roelcke. (Also Salmonella Typ Holstein, Roelcke, Cent. f. Bakt., I Abt., Orig., 137, 1936, 464.) According to Kauffmann (Ztschr. f. Hyg., 119, 1937, 352) the 0-antigens of this rapid fermenter of salicin and weak indole-former are identical with those of Salmonella poona. The H-antigens have not been compared as yet. Salmonella icteroides (Sanarelli) Ber- gey et al. (Bacillo icteroide, Sanarelli, II Policlinico, 4, 1897, 412; Bacillus icteroides Sanarelli, British Med. Jour., July 3, 1897, 7; Bacterium icteroides Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 241; Bergey et al., Manual, 1st ed., 1923, 218.) From yel- low fever cadavers. See Manual, 5th ed., 1939, 604 for a description of this species. Salmonella iwo-jima Lindberg and Bayliss. (Jour. Inf. Dis., 79, 1946, 92.) Isolated from a soldier on Iwo- Jima during a routine examination of food handlers. Belongs to Group C. Antigenic structure: VI, VIII: i: 1, 5 . . . Described too recently to be included in the main bodj' of the text. Salmonella liceagi Leon. (Rev. Inst. Salubridad y Enferm. Trop., 3, 1942, 273.) From feces. This probably be- longs in the coliform group. Salmonella macfadyeanii (Weldin and Levine) Weldin. {Bacterium macfad- yeanii Weldin and Levine, Abst. Bact., 7, 1923, 13; Weldin, Iowa State Jour. Sci., 1, 1927, 168.) Associated with hog cholera. Salmonella -mexicana ^'arela and Olarte. (Rev. Inst. Salubridad y En- ferm. Trop., 4, 1943, 313.) From feces. Salmonella monshaui Carlquist and Coates. (Jour. Bact., 53, 1947, 249.) Isolated from stump of a soldier who suffered traumatic amputation of a leg in the fighting around Monshau, Ger- 532 MANUAL OF DETERMINATIVE BACTERIOLOGY many. Belongs to Group F. Antigenic structure: XXXV: m. t.: — Described too recently to be included in the main body of the text. Salmonella nocardi Pacheco. (Compt. rend. Soc. Biol., Paris, 106, 1931, 372 and 1018.) Pathogenic for parrots and pigeons. Salmonella oahu Lindberg and Bayliss. (Jour. Inf. Dis., 79, 1946, 92.) Isolated from a case of gastroenteritis in a soldier hospitalized on Oahu. Belongs to Group B. Antigenic structure: IV, V, XII: 1, v: 1, 2, 3 . . . Described too recently to be included in the main body of the text. Salmonella ostrei (Besson and Ehr- inger) Hauduroy et al. {Bacillus ostrei Besson and Ehringer, Compt. rend. Soc. Biol., Paris, 87, 1922, 1017; Hauduroy et al., Diet. d. Bact. Path., 1937, 460.) Isolated from oysters. Not pathogenic for laboratory animals. Salmonella para-asiatica (Castellani) Hauduroy et al . ( Bacillus paraasiaticus Castellani, 1916; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 950; Hauduroy et al.. Diet. d. Bact. Path., 1937, 461.) Salmonella para-coagulans (Castellani) Hauduroy et al. {Bacillus para-coagu- lans Castellani, 1914; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 950; Hauduroy et al., Diet. d. Bact. Path., 1937, 461.) Salmonella pauloensis Gomes. (Rev. Inst. Adolf o Lutz, 2, 1942, 231.) May be the same as Salmonella columbensis . Salmonella pseudo-asialica (Castellani) Castellani and Chalmers. {Bacillus pseudo-asialicus Castellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 266; Cas- tellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 940.) Salmonella pseudo-asiatica var. mo- hilis Hauduroy et al. {Bacillus pseudo- asiaticus mohilis Castellani, see Castel- lani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 952; Hauduroy et al.. Diet. d. Bact. Path., 1937, 463.) Salmonella (?) pseudo -Carolina (Cas- tellani) Hauduroy et al. {Bacillus pseu- docarolinus Castellani, 1917; see Cas- tellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 952; Hauduroy et al.. Diet, d. Bact. Path., 1937, 463.) Salmonella {?) pseudo -columbensis (Castellani) Hauduroy et al. {Bacillus pseudo-columbensis Castellani, see Cas- tellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 954; Hauduroy et al.. Diet. d. Bact. Path., 1937, 464.) Salmonella pseudo-morganii (Castel- lani) Hauduroy et al. {Bacillus pseudo- morgani Castellani, see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 954; Hauduroy et al.. Diet. d. Bact. Path., 1937, 464.) Salmonella ranicida Hauduroy et al. (Bacille pathogene isol6 des grenouilles, Gheorghiu and Balmus, Compt. rend. Soc. Biol., Paris, 108, 1931, 1002; Hau- duroy et al., Diet. d. Bact. Path., 1937, 466.) Pathogenic for frogs. Salmonella saipan Lindberg and Bay- liss. (Jour. Inf. Dis., 79, 1946, 92.) Isolated from a case of gastroenteritis in a soldier hospitalized on Saipan. Belongs to Group E. Antigenic struc- ture: III, X, XXVI: Z6. . . . Described too recently to be included in the main body of the text. Salmonella schottmillleri var. alvei Hauduroy et al. {Bacillus paratyphi alvei Bahr, Skand. Veterin. Tidsk., 9, 1919; Hauduroy et al.. Diet. d. Bact. Path., 1937, 469.) Pathogenic for bees and wasps. Salmonella veboda (Castellani) Castel- lani and Chalmers. {Bacillus veboda Castellani, Jour. Trop. Med. and Hyg., 20, 1917, 181; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Bacterium veboda Weldin and Levine, Abst. Bact., 7, 1923, 13.) Salmonella tvatareka (Castellani) Ber- gey et al . ( Bacillus watareka Castellani , Rept. Advisory Committee for Trop. Dis. Research Fund for 1912, London, 1913; Bacterium watareka Weldin and Levine, Abst. Bact., 7, 1923, 13; Bergey et al.. Manual, 1st ed., 1923, 219.) FAMILY ENTEROBACTERIACEAE 533 Salmonella werahensis (Castellani) Hauduroy et al. (Bacillus werahensis Castellani, see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 956; Hauduroy et al.. Diet. d. Bact. Path., 1937, 471.) Salmonella ivesenhergoides (Castellani) Hauduroy et al. (Bacillus ivesenher- goides Castellani, 1916; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 935; Hauduroy et al.. Diet. d. Bact. Path., 1937, 471.) Salmonella willegoda (Castellani) Cas- tellani and Chalmers. (Bacillus wille- goda Castellani; Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 939.) Salmonella woliniae (Castellani) Cas- tellani and Chalmers. (Bacillus woli- niae Castellani, Jour. Trop. Med. and Hyg., £0, 1917, .181; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 939; Bacterium woliniae Weldin and Levine, Abst. Bact., 7, 1923, 13.) Appendix II: The following species have been thought to belong to the genus Eberlhella, i.e., do not produce gas from glucose. Descriptions of nearly all of the species listed in the genus Eberlhella will be found in the Manual, 5th ed., 1939,464-^69. Bacillus subentericus Ford. (Studies from the Royal Victoria Hosp., Mon- treal, ;, 1903, 40; also see Jour. Med. Res., 1, 1901, 218.) From feces. Bacterium typhi flavum Dresel and Stickl. (Deutsche med. Wchnschr., 5^, 1928, 517.) From feces of persons with typhoid fever. Cruickshank (Jour. Hyg., S5, 1935, 354) reports that a variety of yellow chromogenic saprophytes have been identified as belonging to this species, none of which could be regarded as yellow variants of Salmonella tij- phosa (Zopf) White. They apparently belong in the genus Flavobacterium Bergey et al. Eberlhella alcalifaciens de Salles Gomes. (Ri vista do Inst. Adolf o Lutz, 4, 1944, 191.) From catarrhal feces of an infant. Eberthclla belfastiensis (Weldin and Levine) Bergey et al. (Bacterium coli anaerogenes Lembke, Arch. f. Hyg., 26, 1896, 299; Bacterium lembkci xMigula, Syst. d. Bakt., 2, 1900, 417; Bacterium anaerogenes Chester, Man. Determ. Bact., 1901, 135; Bacillus belfastiensis II, Wilson, Jour. Hyg., 8, 1908,543; Bacillus anaerogenes Holland, Jour. Bact., 5, 1920, 217; Bacterium belfastiensis Weldin and Levine, Abst. Bact., 7, 1923, 13; Bergey et al., Manual, 1st ed., 1923, 226; Bacillus coli anaerogenes Kerrin, Jour. Hyg., 2S, 1928, 4 ; Escherichia anaerogenes Bergey et al.. Manual, 3rd ed., 1930, 321; Castellanus colianaerogenes Castellani, Cent. f. Bakt., I Abt., Orig., 125, 1932, 42.) From feces. Eberlhella bentotensis (Castellani and Chalmers) Bergey et al. (Bacillus bento- tensis Castellani, Cent. f. Bakt., I Abt., Ov'ig., 65, 1912, 262; Bacterium bentotensis Weldin and Levine, Abst. Bact., 7, 1923, 15; Bergey et al.. Manual, 1st ed., 1923, 227; Castellanus bentotensis Castellani, Cent. f. Bakt., I Abt., Orig., 125, 1932, 42.) From the intestinal canal. Eberlhella chylogena (Ford) Bergey et al. (Bacillus chylogenes Ford, Stud- ies from the Royal Victoria Hospital, Montreal, 1, No. 5, 1903, 62; Bergey et al.. Manual, 1st ed., 1923, 224.) From the intestinal canal. Eberlhella dubia (Chester) Bergey et al. (Meiner Bakterie, Bleisch, Ztschr. f. Hyg., 13, 1893, 31; Bacillus dubius Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 323; Bacillus bleischii Kruse, ibid., 704; Bacterium dubius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., S>, 1897, 93; Bergey et al.. Manual, 1st ed., 1923, 225.) From the intestinal canal. Eberlhella enterica (Ford) Bergey et al. (Bacillus enter icus Ford, Studies from the Royal Victoria Hospital, Montreal, /, No. 5, 1903, 40; also see Jour. Med. Research, /, 1901, 211; not Bacillus 534 MANUAL OF DP:TERMINATIVE BACTERIOLOGY entericus Castellani, 1907 (Enteroides enter icus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 941); Bergey et al., Manual, 1st ed., 1923, 223.) From the intestinal canal. Eberthella insecticola Steinhaus. (Jour. Bact., 4^, 1941, 762 and 769.) From the intestinal tracts of grass- hoppers, milkweed bugs and stinkbugs. Eberthella kandiensis (Castellani) Ber- gey et al. (Bacillus kandiensis Castel- lani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 262; Eberthus kandiensis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 936; Bacterium kandiensis Weldin and Levine, Abst. Bact., 7, 1923, 13; Bergey et al., Manual, 1st ed., 1923, 225.) From feces. Eberthella lewisii Weldin. (Organism B32, Lewis, Local Gov. Board Rept. Med. Suppl. London, 1910-]!, Appen. B, No. 2, 1911, 314; Bacterium lewisii Wel- din and Levine, Bact. Abst., 7, 1923, 13; Weldin, Iowa State Col. Jour. Sci., 1, 1926, 172.) From feces of a normal child. Eberthella oedematiens Assis. (Boletin do List. Vital, Brazil, 5, 1928.) From the intestinal canal. Eberthella uxijphila (Ford) Bergey et al. (Bacterium oxyphilum Ford, Studies from the Royal Victoria Hos])i- tal, Montreal, ;, Xo. 5, 1903, 49; Bergey etal.. Manual, 1st ed., 1923,224.) From the intestinal canal. Eberthella paulocnsis Mello. (Jornal dos Clinicos, Rio de Janeiro, Xo. 18-30, Sept., 1937, 7 pp.) From feces of a dysentery patient . Eberthella priztnitzi (Cr.stellani and Chalmers) Hauduroy et al. (Bacillus priztnitzi Castellani, .Jour. Trop. Med. and Hyg., 20, 1917, 182; Eberthus prizt- nitzi Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 936; Bac- terium priztnitzi Weldin and Levine, Abst. Bact., 7, 1923, 13; Hauduroy ot al.. Diet. d. Bact. Path., 1937, 186.) From cases of paraenteric fever. Eberthella proteosimilis Wassilien. (Cent. f. Bakt., 1 Abt., Orig., 151, 1944, 423.) Colonies show motility on agar. From feces of a dysentery patient. Eberthella pyogenes (Migula) Bergey et al. (Bacillus pyogenes foetidus Pas- set, Fortschr. der Med., 1885; Bacillus foetidus Trevisan, I generi e le specie delle Batteriacee, 1889, 16; Bacterium pyogenes foetidus Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 141; Bacterium pyogenes Migula, Syst. der Bakt., 3, 1900, 381; Lankoides pyogenes Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 938; Bacillus pyo- genes-foetidus Holland, Jour. Bact., 5, 1920, 220; Bergey et al.. Manual, 1st ed., 1923, 226; Castellanus pyogenes Castel- lani, Cent. f. Bakt., I Abt., Orig., 125, 1932, 42.) From a rectal abscess. Eberthella talarensis (Castellani) Ber- gey et al. (Bacillus talavensis Castel- lani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 262; Eberthus talavensis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 936; Bacterium talavensis Weldin and Levine, Abst. Bact., 7, 1923, 13; Bergey et al., Manual, 1st ed., 1923, 225.) From the intestinal canal. Eberthella tarda Assis. (Boletin do Inst. Vital, Brazil, 5, 1928.) From the intestinal canal. Eberthella wesenbergi (Castellani and Chalmers) Hauduroy et al. (Bacillus iresenberg Castellani, 1913; Weseribergus ivesenbergi Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 940; Hauduroy et al.. Diet. d. Bact. Path., 1937, 191.) Eberthella ivilsonii Weldin. (Bacillus hclfastiensis V, Wilson, Jour. Hyg., 8, 1908, 543; Weldin, Iowa State Col. Jour. Sci., ;, 1926, 174.) From feces. Eberthella xenopa Schrire. (Trans. Iloj-al Soc. So. Africa, 17, 1928, 43.) From wound infection in frogs. Wese7ibergus fcrmentosus Castellani and Chalmers. (Man. Trop. Med., 3rd ed., 1919, 940.) From blood. Isolated by Archibald in the Anglo-Egyptian Sudan. FAMILY EXTEKOBACTERIACEAi: 535 Genus II. Shigella Castellani and Chalmers* (Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 936; subgenera, Flex- nerella and Shigella, Castellani and Chalmers, ibid., 938; Castellanus Carruti, Jour. Trop. Med. and Hyg., July 15, 1930; Proshigella Borman, Stuart and Wheeler, Jour. Bact., Jt8, 1944, 363.) Xamed for Prof. I. Shiga, the Japanese bacteriologist who dis- covered the dysenterj- bacillus in 1898. Non-motile rods, although cultures of some of the less well-known species have been reported as motile. Produce acid but no gas from carbohydrates except with some types of Shigella paradysenteriae. Do not liquefy gelatin. Some species produce acid from lactose and form indole. Some species reduce trimethylamine oxide to trimethylamine, others do not.f Some species will grow at 45.5°C (Eijkman test).t Pathogenic (causing dysenteries) or non-pathogenic species, all living in the bodies of warm-blooded animals. Carried by polluted water supplies and by flies. The type species is Shigella dysenteriae (Shiga) Castellani and Chalmers. Key to the species of genus Shigella.*'' I. Xo acid from mannitol. A. Xo acid from lactose. IMilk not coagulated. 1. Indole not produced. a. Acid but no gas from glucose. 1. Shigella dysenteriae. aa. Acid and a small amount of gas from glucose. 4a. See Shigella paradysenteriae (Type Xewcastle). 2. Indole produced. 2. Shigella ambigua. B. Acid formed slowlv from lactose. 1. Indole not produced. 3. Shigella gintottensis. II. Acid from mannitol (one type produces a small amount of gas). A. Xo acid from lactose. 1. Xo acid from rhamnose, .xylose or dulcitol. 4. Shigella paradysenteriae. 2. Acid from rhamnose, xylose and dulcitol. 5. Shigella alkalescens. 3. .\cid from x.vlosc but not from dulcitol. 6. Shigella pfaffii. B. Acid formed slowly from lactose. 1. Indole not produced. a. Acid from rhamnose. Xone from .xylose. 7. Shigella sonnei. aa. Xo acid from rhamnose. Acid from xylose. 8. Shigella cquirulis. * Completely revised by Dr. Frederick Smith, McGill University, Montreal, P. Q., Canada, December, 1938; further revision, April, 1946. t Wood, Baird and Keeping, Jour. Bact., 46, 1943, 106. t Stuart and Rustigian, Jour. Bact., 46, 1943, 105. ** See Weil, Jour. Immunology, 55, 1947, .363-405. 536 MANUAL OF DETERMINATIVE BACTERIOLOGY 2. Indole produced. a. Acid from dulcitol. aa. No acid from dulcitol. III. Action on mannitol unknown. A. No acid from lactose. 1. Indole is produced. 1. Shigella dysenteriae (Shiga) Cas- tellani and Chalmers. (Bacillus of Jap- anese dysentery, Shiga, Cent. f. Bakt., I Abt., 23, 1S98, 599; Bacillus dysenteriae Shiga, Cent. f. Bakt., I Abt., 24, 1898, 817; Bacillus japaniciis Migula, Syst. d. Bakt., 2, 1900, 755; Bacillus shigae Chester, Man. Determ. Bact., 1901, 228 Bacillus dysentericus Ruffer and Will more, Brit. Med. Jour., 2, 1909, 862 Bacterium dysenteriae Lehmann and Neumann,Bakt.Diag.,5Aufl.,,?, 1912,348 not Bacterium dysenteriae Chester, Man Determ. Bact., 1901, 145; Castellani and Chalmers, Man. Trop. Med., 3rd ed. 1919, 935; Bacterium shigae Holland Jour. Bact., 5, 1920, 220; Eberthella dysenteriae Bergej^ et al.. Manual, 2nd ed., 1925, 250.) Latinized, of dysentery. Rods: 0.4 to 0.6 by 1.0 to 3.0 microns, occurring singly. Non-motile. Gram- negative. Gelatin colonies: Small, grayish, smooth, homogeneous, entire to slightly undulate. Gelatin stab: Grayish surface growth. No liquefaction. Agar slant : Grayish, filiform to echinu- late, smooth, entire to undulate growth. Broth: Slightly turbid, with grayish sediment. v Litmus milk: Slightly acid, then alka- line. Potato: Delicate, grayisli io slightly brownish streak. Indole not produced. Nitrites produced from nitrates. Acid but no gas from glucose, fructose, raffinose, glycerol and adonitol. Does not attack arabinose, xylose, maltose. 9. Shigella ceylonensis. 10. Shigella madampcnsis. 11. Shigella seplicemiae. lactose, sucrose, salicin, mannitol, dul- citol or rhamnose. Does not reduce trimethylamine oxide (Wood et al.. Jour. Bact., 46, 1943, 106). Aerobic, facultative. Optimum temperature 37°C. Does not grow at45.5°C (Eijkman's reaction, Stuart et al.. Jour. Bact., 46, 1943, 105). Serologically homogeneous and differ- ent from the other species of Shigella. Forms a potent exotoxin. Source: From widespread epidemics of dysentery in Japan. Habitat : A cause of dysentery in man and monkeys. 2. Shigella ambigua (Andrewes) Wei-' din. (Bazillus Schmitz, Schmitz, Ztschr. f. Hyg., 84, 1917, 449; Bacillus ambiguus Andrewes, The Lancet, 194, 191S,5&0; Bacillus dysenteriae' 'Schmitz", Murray, Jour. Roy. Army Med. Corps, 3/, 1918,257; Bacterium ambiguum Levine, Abst. Bact., 4, 1920, 15; not Bacterium ambiguum Chester, Del. Col. Agr. Exp. Sta. Ann. Rept., 11, 1900, 59; Eberthella ambigua Bergey et al.. Man- ual, 1st ed., 1923, 229; Bacillus para- dysenteriae X, Stutzer, Cent. f. Bakt., I Abt., Orig., 90, 1923, 12; Bacterium schmitzil Weldin and Levine, Abst. Bact., 7, 1923, 13; Weldin, Iowa State College Jour. Sci., 1, 1927, 177; Shigella schmitzii Ilauduroy et al.. Diet. d. Bact. Path., 1937, 496.) From Latin, uncertain. Morphology and colony characters in- distinguishable from those of Shigella dysenteriae. Acid from glucose and rhamnose. FAMILY ENTEROBACTERIACEAE 537 Does not attack xylose, maltose, lactose, sucrose, dextrin, glycerol, mannitol or dulcitol. Indole is produced. Does not reduce trimethylamine oxide (Wood et al., Jour. Bact., 46, 1943, 106). Aerobic, facultative. Optimum temperature 37°C. Does not grow at 45.5°C (Stuart et al., Jour. Bact., J^6, 1943, 105). Serologicall}" homogeneous and differ- ent from the other species of Shigella. Does not form an exotoxin. Source : Found in feces in a dysentery epidemic in a prison in Germany. Habitat: A cause of human dysentery. 3. Shigella gintottensis (Castellani) Hauduroy et al. (Bacillus ginlollensis Castellani, 1910; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 948; Lankoides gintottensis Castel- lani and Chalmers, ibid. ,938; Castellanus gintottensis Castellani, 1930; Castellani, Jour. Trop. Med. and Hyg., 36, 1933, 109; Hauduroy et al., Diet. d. Bact. Path., 1937, 488.) Rods: Non-motile. Gram-negative. Morphology and cultural characters indistinguishable from those of Shigella dysenteriae. Litmus milk: Acid and coagulation; decolorized. Indole not formed. Acid, but no gas, from lactose, glu- cose, arabinose and galactose. No acid from sucrose, dulcitol, mannitol, mal- tose, dextrin, raffinose, adonitol, inulin, sorbitol, levulose, inositol, salicin and glycerol. Antigenic structure not known. Source: From feces in cases of dysen- tery. Habitat : A cause of human dysentery. 4. Shigella paradysenteriae (Collins) Weldin. {Bacillus dysenteriae Flexner, Phil. Med. Jour., 6, 1900, 414; Bacillus dysenteriae Hiss and Russell, Medical News, 82, 1903, 289; Bacillus dysenteriae Strong, Jour. Amer. Med. Assoc, 35, 190G, 498; Bacillus paradysenteriae Col- lins, Jour. Inf. Dis., 2, 1905, 620; includes weakly toxic strains of dysentery bacilli, Groups I and II, Sonne, Cent. f. Bakt., I Abt., Orig., 75, 1915, 408; Shigella flexneri Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 937; Shigella dysenteriae (Hiss and Russell, and Strong types) Castellani and Chalmers, ihid., 937; not Shigella paradysenteriae Castel- lani and Chalmers, ibid., 937; Bacillus flexneri Levine, Jour. Inf. Dis., 27, 1920, 31; Bacterium flexneri Levine, Abst. Bact., 4, 1920, 15; Bacterium dysenteriae (Flexner type) and Bacterium paradysen- teriae Holland, Jour. Bact., 5, 1920, 215; Eberthella flexneri Weldin and Levine, Abst. Bact., 7, 1923, 13; Eberthella para- dysenteriae Bergey et al., Manual, 1st ed., 1923, 230; Weldin, Iowa State Col- lege Jour. Sci., 1, 1927, 178.) Latinized, like dysenter^^ Rods: 0.5 by 1.0 to 1.5 microns. Non- motile. Gram-negative. Morphologically these organisms are like Shigella dysenteriae. Culturallj^ these organisms differ from Shigella dysenteriae in that they ferment mannitol. No acid is produced from lactose, rhanmose, xylose or dulcitol. Does not reduce trimethylamine oxide (Wood et al.. Jour. Bact., 46, 1943, 106). Does not form a potent exotoxin. Aerobic, facultative. Optimum temperature 37 °C. Does not grow at 45.5°C (Stuart et al., Jour. Bact., 46, 1943, 105). Antigenically the organisms of this species are not homogeneous. Boyd (Trans. Roy. Soc. Trop. Med. and Hyg., 33, 1940, 553) has shown that the mannitol -fermenting Shigella include many organisms previously unknown or unclassified because they did not agree with the classical types of Andrewes and Inman (Med. Res. Council, Special Rept. Ser. No. 42, London, 1919). With these, on grounds of antigenic structure, will be included the gas-forming Man- chester bacillus of Downie, Wade and Young (Jour. Hyg., 33, 1933, 196) and 538 Manual of determinative bacteriology both maliliitol-fermeutiiig and non-fer- menting Newcastle bacilli (Clayton and Warren, Jour. Hyg., 28, 1929, 355 and 29, 1929, 191). The following tables are taken from Boyd (loc. cit.). Table 1. — Classification of Shigella parody senteriae. New Name Bacillus dysenteriae Flexner I Bacillus dysenteriae Flexner II Bacillus dysenteriae Flexner III Bacillus dysenteriae Flexner IV Bacillus dysenteriae F'lexner V Bacillus dysenteriae Flexner VI Bacillus dysenteriae Boyd I Bacillus dysenteriae Boyd II Bacillus dysenteriae Boyd III Old Name Andrewes Inman (Flexner) Andrewes Inman (Strong) Andrewes Inman Z Type 103 Type P 119 88-Newcastle- Manchester group Type 170 Type P 2S8 Type Dl and V and The six Flexner types possess a com- mon group antigen and separate type- specific antigens. The three Boyd types are distinct antigenically from each other and from the Flexner types. Two new Flexner types (Type 953 = provisional Type VII and Type 1296/7 = provisional Tj^pe VIII) have been described by Francis (Jour. Path, and Bact., 58, 1946, 320) as this section goes to press. Also see Boyd (ibid., 297). Table 2. — Subclassification of Bacillus dysenteriae Flexner VI (including the Newcastle bacillus). Type 88 (33 per cent of strains) Type 88 (66 per cent of strains) Manchester ba- cillus Newcastle bacil- lus o o 3 O 'S c - A A - A A - AG AG - AG - Dulcitol (late) A (late) AG (late) AG Source; From feces in cases of dysen- tery. Habitat: A cause of dysentery in man. A cause of summer diarrhoea in children. Note: The term Bacillus paradysen- teriae is used by Kruse (Mtinch. med. Wchnschr., 1917, 1309) for the Esche- richia coZ/-like motile and gas-forming Gram-negative rods that have been found to cause dysentery-like diseases. Kruse (Deut. med. Wchnschr., 27, 1901, 388) uses the term pseudodysentery for the group that includes the Flexner, Strong, and Hiss and Russell types. See Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 456. Gardner (Med. Res. Council, System of Bacteriol- ogy, 4, 1929, 170) states that "Kruse's terms B. dysenteriae for Shiga, and Ba- cillus pseudodysenteriae for the Flexner- Sonne-Schmitz groups have, however, never taken root outside the German- speaking world". 4a. Shigella paradysenteriae (Type Newcastle). (Clayton and Warren, Jour. Hyg., 28, 1929, 355 and 29, 1929, 191; Bacillus dysenteriae Fle.xner VI in part, Boyd, Trans. Roy. Soc. Trop. Med. and Hyg., 33, 1940, 553.) Rods: Non-motile. Gram-negative. In peptone water solution, lactose, mannitol, and sucrose not fermented. Glucose, maltose and dulcitol fermented. Peculiarities of the organism are: (1) Occasionally a slight bubble of gas is produced from glucose and dulcitol, (2) when the substrate is dissolved in beef extract broth, glucose, dulcitol and mal- tose are always fermented to gas and acid. Does not reduce trimethylamine oxide (Wood et al., Jour. Bact., 46, 1943, 106). Optimum temperature 37 °C. Does not grow at 45.5°C (Stuart et al.. Jour. Bact., 46, 1943, 105). Aerobic, facultative. Serologically related to the mannitol- fermenting strains of Shigella paradysen- teriae. FAMILY ENTEROBACTERIACEAE 539 Source : Isolated in 1925 from a case of diarrhoea in Xewcastle-on-Tyne, Eng- land . Habitat : A cause of human dysentery. 4b. Shigella paradysenteriae (Tj'pe Manchester). (Downie, Wade and Young, Jour. Hyg., 33, 1933, 196; Bacil- lus dysenteriae Flexner VI in part, Boj^d, Trans. Roy. Soc. Trop. Med. and Hyg., 33, 1940, 553.) Characters as for Type Xewcastle ex- cept that acid and gas are produced from mannitol. Does not produce gas from maltose. Serologically related to the non-manni- tol-fermenting strains of Shigella para- dysenteriae. Source : Five strains were isolated from cases of dysentery at Denton near ^Nlan- chester, England. One strain came from a case of dysentery in Nigeria. Habitat: A cause of human dysentery. 5. Shigella alkalescens (Andrewes) Weldin. {Bacillus alkalescens An- drewes, The Lancet, London, 19i, 1918, 560; Bacterium alkalescens Levine, Jour. Inf. Dis., 27, 1920, 31; Eherthella alka- lescens Bergey et al.. Manual, 1st ed., 1923, 231; Weldin, Iowa State College Jour. Sci., /, 1927, 179; Proshigella alka- lescens Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 363.) From the chemical term, alkaline. Rods: 0.5 by 1.0 to 1.5 microns, occur- ring singly and in pairs. Non -motile. Gram-negative. Gelatin stab: Xo liquefaction. Agar slant: Abundant, transparent, often iridescent growth. Broth: Turbid. Litmus milk: Acid, then alkaline. Potato: Moderate, grayish growth. Indole is formed. Acid but no gas from glucose, xjdose, rhamnose, maltose, mannitol and dulci- tol. Sucrose is fermented by some strains. Does not attack lactose, dex- trin or salicin. Reduces trimethylamine oxide to tri- methylamine (Wood et al., Jour. Bact., 46, 1943, 106). In contrast to all other species of the genus, will also produce trimethylamine from choline (Wood and Keeping, Jour. Bact., 47, 1944, 309). Aerobic, facultative. Optimum temperature 37°C. Grows at 45.5°C (Eijkman's reaction, Stuart et al., Jour. Bact., 46, 1943, 105). Not pathogenic. Not agglutinated by Shiga immune serum. Source: From feces in cases of dysen- tery. Habitat: Intestinal canal. 6. Shigella pfaffii (Hadley etal.) Wel- din. (Bacillus der kanarienvogelseuche, Pfaff, Cent. f. Bakt., I Abt., Orig., 38, 1905, 276; Bacterium pfaffi Hadley, El- kins and Caldwell, Rhode Island Agr. E.xp. Sta. Bull. 174, 191S, 169; Bacillus pfaffi Hadley, Elkins and Caldwell, ibid.. 204; Eherthella pfaffi Bergey et al.. Manual, 1st ed., 1923, 232; Weldin, Iowa State College Jour. Sci., 1, 1927, 180.) Named for Dr. Franz Pfaff of Prague who isolated this species. Description largely from Hadley et al. {loc. cit., 180). Rods: 0.5 by 1.0 to 2.0 microns, occur- ring singly. Non-motile. Gram-nega- tive. Gelatin colonies : Small, grayish, trans- lucent. Gelatin stab: No liquefaction. Agar colonies: Small, yellowish-gray, homogeneous, translucent, entire. No odor. Agar slant: Slight, yellowish-gray, translucent streak. Broth: Turbid, with flocculent sedi- ment (Pfaff, loc. cit., 280). Litmus milk: Unchanged. Potato: Moderate, whitish streak. Acid but no gas from glucose, fructose, arabinose, xylose, maltose, dextrin, salicin and mannitol. Does not attack lactose, sucrose, raffinose, inulin, adoni- tol or dulcitol. Indole not formefl. No hydrogen sulfide produced. 540 MANUAL OF DETERMINATIVE BACTERIOLOGY Nitrites not produced from nitrates. Pathogenic for canaries, sparrows, pigeons, white mice, guinea pigs and rabbits. Not pathogenic for chiclcens (Pfaff, loc. cit., 280). Aerobic, facultative. Optimum temperature 37 °C. Source: First encountered in an epi- demic of septicemia in canaries. Caused a necrotic enteritis. Habitat: Not known from other sources. 7. Shigella sonnei (Levine) Weldin. (Duval's bacillus, Duval, Jour. Amer. Med. Assn., 43, 1904, 381; Pseudodysen- tery bacillus E, Kruse, Deutsche med. Wchnschr., 23, 1907, 292, 338; Bacillus ceylonensis A, Castellani, Jour. Hyg., 7, 1907, 1; Group III of Sonne, Sonne, Cent. f. Bakt., I Abt., Orig., 75, 1915, 408; Bacillus dispar (in part) Andrewes, Lancet, 1, 1918, 560 (see Shigella cey- lonensis and Shigella madampensis) ; Ba- cillus of Sonne, Thj0tta, Jour. Bact., 4, 1919, 355; Bacterium sonnei Levine, Jour. Inf. Dis., 27, 1920, 31; Bacillus dysenteriae Sonne, Smith, Jour. Hyg., 23, 1924, 94; Weldin, Iowa Sta. Coll. Jour. Sci., /, 1927, 182; Castellamis kruse- castellani Cerruti, Jour. Trop. Med. and Hyg., 33, 1930, 207; Shigella paradysen- teriae var. sonnei Bergey et al., INIanual, 4th ed., 1934, 393; Proshigella sonnei Borman, Stuart and Wheeler, Jour. Bact., 48, 1944, 363.) Named for Dr. Carl Sonne, who worked with this organism. Rods: Non -motile. Gram-negative. Gelatin: No liquefaction. Agar colonies: Cultures dissociable into two types: (1) Glistening sur- face, 2 mm in diameter in 24 hours, soft, grayish, edge entire; (2) Granular sur- face, 3 to 4 mm in diameter in 24 hours, soft, grayish, edge tending to spread un- evenly, surface developing, after some days, papillae (daughter colonies) which are lactose-fermenting. Some colonies of type 1 change to type 2 on continued incubation. The colony types do not breed true. Broth: Many authors stress the floccu- lent growth, associated with spontaneous agglutination in saline solution. These appear to be dependent on growth condi- tions and time of incubation. Litmus milk: Acid and with about 50 per cent of strains coagulation. Coagu- lation tends to occur later than the fer- mentation of lactose in peptone water. Indole is not produced. Acid, but no gas, from lactose (about 2 per cent of strains are lactose -negative after 2 months incubation), glucose, fructose, maltose, galactose, rhamnose, mannitol, arabinose, raffinese and suc- rose. No acid from dulcitol, inulin, inositol, adonitol, xylose (xylose is occa- sionally fermented) and salicin. Fermentation of substances other than the monosaccharides may require days or weeks. Reduces trimeth3damine oxide to tri- methylamine (Wood et al.. Jour. Bact., 46, 1943, 106). Serologically Shigella sonnei is divis- ible into two types, which do not corre- spond with the colony types described above. Most freshly isolated strains absorb agglutinins completely from all Shigella sonnei antisera, while most stock strains absorb only partially from other than antisera of the second serological type. There exist minor serological re- lationships between Shigella sonnei and Shigella paradysenteriae, Shigella alka- lescens and Shigella madampensis. Optimum temperature 37 °C. Grows at 45.5°C (Stuart et al., Jour. Bact., 46, 1943, 105). Source: From feces in cases of dysen- tery. Habitat: A cause of mild dysentery in man; summer diarrhoea in children. 8. Shigella equirulis (de Blieck and van Heelsbergen) Edwards. (Bacillus nephritidis equi Meyer, Transvaal Dept. Agr. Rept. Gov. Bac, 1908-1909, 122; Bacterium viscosum equi Magnusson, FAMILY ENTEROBACTERIACEAE 541 Svensk. Veterinartijdskr., 1917, 81 ; also see Jour. Comp. Path, and Therap., 32, 1919, 143; Bacillus equuli van Straaten, Verslag van den Werkzaamheden der Rijksseruminrichting voor 1916-1917, Rotterdam, 1918, 75; Bacterium pyosep- Hcus equi de Blieck and van Heelsbergen, Tydschr. v. Diergeneesk., J^6, 1919, 492; Bacillus equirulis de Blieck and van Heelsbergen, ibid., 496; Bacterium pyo- septicum viscosum IMeissner, Deut. tier- artzl. Wchnschr., 29, 1921, 185; Bacte- riiim pyosepiicum (viscosmn) equi Ltitje, Deut. tierarztl. Wchnschr., 29, 1921, 463; Bacterium pyosepticum (viscosum) Meiss- ner and Berge, Deut. tierarztl. Wchn- schr., 30, 1922, 473; Bacterium pyosepti- CMmMeissner, Deut. tierartzl. Wchnschr., 31, 1923, 348; Bacterium pyosepticum equi Landien, Inaug. Diss., Hanover, 1923; Bacillus pyosepticus Clarenberg, Ztschr. f. Infektskr. u. Hyg. d. Haust., 27, 1924, 193; Bacterium equi Weldin and Levine, Abst. Bact., 7, 1923, 13; Eberthella vis- cosa Snyder, Jour. Amer. Vet. Med. Assoc, b'6, 1925, 481; Shigella equi Wel- din, Iowa Sta. Col. Jour. Sci., 1, 1927, 121; Shigella viscosa Bergey et al., Manual, 3rd ed., 1930, 363; Edwards, Kentucky Agr. Exp. Sta. Res. Bui. 320, 1931.) While awaiting further information, the binomial introduced by Edwards is used for this species although Haupt of Leipzig points out in a personal com- munication to Edwards (1934) that van Straaten's original name was Bacillus equuli. The binomial Bacillus equirulis is stated to have appeared first in the article by de Blieck and van Heels- bergen, loc. cit. Description from Edwards {loc. cit.). Rods: 0.3 to 0.4 by 0.4 to 0.8 micron, occurring singly, in chains and filaments. Young cultures (8 to 10 hrs.) frequently show long filaments and streptococcus- like chains as well as large, yeast-like bodies with projections. Rough mucoid colonies consist of short, oval rods. Smooth colonies contain long filaments and streptococcus-like chains. Rough colonies are always mucoid. Non-mu- coid colonies are alwaj's smooth. Cap- sules described but uncertain. Non- motile. Gram-negative. Gelatin colonies: Grayish -white, cir- cular, translucent. Gelatin stab: Nail-head, moderate growth along line of stab. No lique- faction. Agar colonies: 3 to 5 mm at 48 hours. Semi-solid, tough, adherent, circular, grayish-white, smooth, moist, glisten- ing. Rough variants and dwarf colonies. Agar slant: Grayish-white, viscid growth, covering the surface. Viable 8 to 10 days. Broth: Masses form on side of tube. At times a thin gra^dsh pellicle. Gray- ish, tough, ropy sediment. Eventually diffuse turbiditj' which is highly viscous. Viabilitj^ 2 to 4 weeks. Litmus milk: Slowly acidified; slimy, viscid. Sometimes coagulation and re- duction. Potato: No visible growth. Indole not formed. Nitrites produced from nitrates. Voges-Proskauer test negative. Acid but no gas from glucose, fructose, xylose, lactose, galactose, maltose, suc- rose, mannitol and raffinose. Dextrin usually fermented. No action in rham- nose, dulcitol, sorbitol or inositol. Usually no action in salicin, adonitol and arabinose. Does not reduce trimethylamine oxide (Wood et al.. Jour. Bact., 46, 1943, 108). Does not grow at 45.5°C (Stuart et al., Jour. Bact., ^6, 1943, 105). Optimum temperature 37°C. Aerobic, facultative. Not pathogenic for small experimental animals. Produces abscesses and stiff- ening of the joints when injected subcu- taneously in horses. Serologically heterogeneous. Nothing is known of its antigenic relations to other members of the genus. Haupt writes in a personal communication that comparative serological studies indicate 542 MANUAL OF DETERMINATIVE BACTERIOLOGY that this species should be placed in the genus Actinohacilhis. Distinctive characters : Differentiation from Shigella sonnei is made on cultural and morphological grounds and imme- diate fermentation of lactose. Source: Isolated from cases of joint- ill in foals. Habitat: Causes joint-ill in foals. 9. Shigella ceylonensis (Castellani) Weldin. {Bacillus ceylonensis B, Cas- tellani, Jour. Hyg., 7, 1907, 1; Bacillus dispar (in part) Andrewcs, Lancet, 1, 1918, 560 (see Shigella madampensis and Shigella sonnei. Andrewes included in Bacillus dispar all lactose -fermenting members of the dysentery group) ; Lankoides ceylonensis B, Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 938; Eberthella dispar Bergey et al., Manual, 1st ed., 1923, 232 (see Shigella madavvpensis) ; Weldin, Iowa Sta. Coll. Jour. Sci., 1, 1927, 182; Castallanus cas- tellanii Cerruti, Jour. Trop. Med. and Hyg., 33, 1930, 207.) Latinized, per- taining to Ceylon. Rods: Non-motile. Gram-negative. Morphology and colony characters indistinguishable from those of Shigella dysenteriae . Gelatin not liquehed. Litmus milk: Acid with coagulation. Indole is formed. Acid, but no gas, from lactose, glu- cose, fructose, sucrose, mannitol, dulci- tol, maltose, .xylose, arabinose, rham- nose, sorbitol, raffinose, de.xtrin and glycerol. Inulin, inositol, adonitol and salicin not fermented (salicin differen- tiates Shigella cetjlonensis from Bac- terium coli anacrogencs Lembke, Arch. f. Hyg., 26, 1896, 299). Substances other than the monosac- charides are characteristically fermented slowly. Reduces trimethylamine oxide to tri- methylamine (Wood et al., Jour. Bact., 1,6, 1943, 106). Pathogenic for guinea pigs anil ral)bits. Serologically the organism is stated by Castellani to be homogeneous and completely different from Shigella mad- ampensis and Shigella sonnei. The rela- tions to other members of the dysenter}' group have not been stated. Optimum temperature 37 °C. Grows at 45.5°C (Stuart et al., Jour. Bact., 46, 1943, 105). Source: Isolated from the stools and intestines of persons suffering from dysentery. Habitat: A cause of dysentery in man. 10. Shigella madampensis (Castellani) Weldin. {Bacillus madampensis Cas- tellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 262; Bacillus dispar (in part) An- drewes, Lancet, 1, 1918, 560 (see Shigella cetjlonensis and Shigella sonnei) ; Lan- koides madampensis Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 938; Bacterium dispar Levine, Abst. Bact., 4, 1920, 15; Eberthella dispar Bergey et al.. Manual, 1st ed., 1923, 232 (see Shigella ceylonensis); Weldin, Iowa Sta. Coll. Jour. Sci., .?, 1927, 181a; Shi- gella dispar Bergey et al., Manual, 3rd ed., 1930, 364; Proshigella dispar Borman, Stuart and Wheeler, Jour. Bact., 4S, 1944, 363.) Xeter (Bact. Rev., 6, 1942, 26) com- bines Shigella ceylonensis and S. madam- pensis into a single species which he names Shigella castellani i. Strains currently existing in various Type Collections as Bacillus dispar have biochemical properties indistinguishable from those described for Shigella madam- pensis (Glynn and Starkey, Jour. Bact., 37, 1939, 315). Rods: Non-motile. Gram-negative. Morphology and colony characters in- distinguishable from those of Shigella (lyscnteriae. Gelatin not liquefied. Indole is formed. Litmus milk: Acid with coagulation. Acid, but no gas, from lactose, mal- tose, sucrose, arabinose, xylose, glycerol, mannitol, rhamnose, glucose, fructose, galactose and dextrin. Dulcitol, salicin, inulin, inositol and adonitol not fer- mented. -FAMILY ENTERCBACTERIACEAE 543 Substances other than monosaccharides are characteristically fermented slowly. Reduces trimethj'lamine oxide to tri- methylamine (Wood et al., Jour. Bact., Jt6, 1943, 106). Serologically the organism is stated by Castellani to be homogeneous and completely different from Shigella ceij- lonensis and Shigella sonnei. According to Andrewes {loc. cit.), Bacillus dispar is serologically distinct from Shigella alkalescens and Shigella paradysenteriae. Fifteen strains (Glynn and Starkey, loc. cit.) from various sources, labelled Bacillus dispar and conforming to the above description, proved to be sero- logically heterogeneous. Optimum temperature 37 °C. Grows at 45.5°C (Stuart et al.. Jour. Bact., 46, 1943, 105). Source: Isolated from human stools and intestines. Habitat: Considered by Castellani to be a cause of colitis and cystitis. 11. Shigella septicaemiae (Bergey et al.) Bergej- et al. (Bacillus septicaemiae anserum exsudativae Riemer, Cent. f. Bakt., I Abt., Orig., 37, 1904, 648; Eber- thella septicaemiae Bergey et al., Manual, 2nd ed., 1925, 250; Bergey et al., Manual, 3rd ed., 1930, 358.) Latinized, of septi- cemia . Small rods: 0.5 by 1.5 to 2.0 microns, occurring singly, in pairs and in threads. Motile. Gram-negative. Gelatin colonies: Small, white, circular. Gelatin stab: Slight, infundibuliform liquefaction, becoming complete in sev- eral weeks. Agar colonies: Circular, transparent, smooth, homogeneous, entire. Agar slant: Soft, grayish-white streak, slightly viscid, becoming transparent. Does not grow on Endo agar. Broth: Slight, uniform turbidity, with slight pellicle formation. Litmus milk: Unchanged. Potato: Xo growth. Blood serum: Yellowish-white streak, the medium becoming brownish and slowly liquefied. Indole is formed after several days. Slight acid and no gas from glucose. Xo acid from lactose. Hydrogen sulfide is formed. X'ot pathogenic for white mice, guinea pigs, chickens or pigeons. ^lildl}^ path- ogenic for ducks. Aerobic. Optimum temperature 37 °C. Source: Isolated from blood, exudates and all of the internal organs of geese. Habitat : Cause of a fatal septicemia in young geese. Appendix: The following species are also found in the literature. Many are incompletely described. Bacillus coli dijsentericum Ciecha- nowski and Nowak. (Cent. f. Bakt., I Abt., Orig., ^3, 1898, 445.) From a case of dysentery. Bacillus dysenteriae Migula. (Bacil- lus of Japanese dysentery, Ogata, Cent, f. Bakt., 11, 1892, 264; Bacillus dysen- teriae liquefaciens Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 2&1; Bacterium dysenteriae liquefaciens Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 102; Migula, Syst. d. Bakt., 2, 1900, 641; not Bacillus dysenteriae Shiga, Cent. f. Bakt., I Abt., 24, 1898, 817; not Bacillus dysenteriae Hiss and Russell, :Medical Xews, 82, 1903, 289; not Bacillus dysenteriae Strong, Jour. Amer. Med. Assoc, 35, 1906, 498; not Bacillus dysen- teriae Sonne, Smith, Jour. Hyg., 23, 1924, 94.) From a case of Japanese dysentery. Motile. Gram-positive. Bacillus dysentericus Trevisan. (Ba- cillus der Dysenteric, Klebs, Cent. f. Bakt., 2, 1887, 248; Trevisan, I generi e le specie delle Batteriacee, 1889, 14; not Bacillus dysentericus Ruffer and Will- more, Brit. Med. Jour., 2, 1909, 862.) From feces. Bacterium pseudodysentericum Kruse. (Kruse, Deutsche med. Wchnschr., 27, 1901, 370, 386; Escherichia pseudodysen- tcriae Bergey et al.. Manual, 1st ed., 1923, 198.) From feces. Motile. Bacterium wakefield Berger. (Jour. 544 MANUAL OF DETERMIXATI\TE BACTERIOLOGY Hyg., U, 1945, 116-119.) From feces. A non-mannitol-fermenting organism of the Flexner group. Wheeler and Stuart (Jour. Bact., 51, 1946, 324) regard this as an anaerogenic paracolon. Shigella albofaciens (Castellani) Hau- duroy et al. (Bacillus albofaciens Cas- tellani, ^Meetings of the Ceylon Branch of the British ]Medical Association, 1905; Hauduroy et al.. Diet, d, Bact. Path., 1937, 482.) Shigella arabinotarda, types A and B, Christensen and Gowen. (Jour. Bact., 47, 1944, 171-176.) From cases of dysentery in U. S. Army in Tunisia. A lactose-negative, mannitol -negative Shigella. Shigella bienstockii (Schroeter) Bergey et al. (Bacillus III, Bienstock, Ztschr. f. klin. Med., 8, 1884; Bacillus coprogenes parvus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 269; Bacillus bienstockii Schroeter, Kryptogamen Flora von Schlesien, 3, 1, 1886, 163; Bacillus parvus Trevisan, I generi e le specie delle Bat- teriacee, 1889, 15; not Bacillus parvus Neide, Cent. f. Bakt., II Abt., 12, 1904, 344; Bacterium coprogenes parvus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Bacterium bienstockii Ches- ter, Man. Deterra. Bact., 1901, 144; Eberthella bienstockii Bergey et al.. Manual, 1st ed., 1923, 227; Bergey et al., Manual , 3rd ed ., 1930 , 360 . ) From feces . Shigella douglasi (Castellani and Chalmers) Hauduroy et al. (Bacillus douglasi Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 946; Hauduroy et al.. Diet. d. Bact. Path., 1937, 484.) Shigella etousae Heller and Wilson. (Jour. Path, and Bact., 58, 1946, 98.) From dysentery outbreak in an army camp in England. Shigella faecaloides (Castellani) Hau- duroy et al. (Bacillus faecaloides Cas- tellani, 1915; see Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 946; Hauduroy et al.. Diet. d. Bact. Path., 1937, 488.) Shigella giumai (Castellani) Hauduroy et al. (Bacillus giumai Castellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 264; Wesenbergiis giumai Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 940; Bacterium giumai Weldin and Levine, Abst. Bact., 7, 1923, 15; Sal- monella giumai Bergey et al., Manual, 1st ed., 1923, 220; Hauduroy et al., Diet. d. Bact. Path., 1937, 488.) Shigella lunavensis (Castellani) Hau- duroy et al. (Bacillus lunavensis Cas- tellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 264; Bacterium lunavensis Weldin and Levine, Abst. Bact., 7, 1923, 16; Hauduroy et al., Diet. d. Bact. Path., 1937, 489.) From feces. Shigella metadysenterica var. A, B, C, and D (Castellani) Hauduroy et al, (Bacillus metadysentericus var. A, B, C, and D, Castellani, 1904; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 946; Dysenteroides metadysen- tericus var. A, B, C, and D, Castellani and Chalmers, Ann. Inst. Past., 34, 1920, 607; Castellanus metadysentericus Cas- tellani, 1930; Hauduroy et al.. Diet. d. Bact. Path., 1937, 489.) From cases of dysentery. Shigela ne^ombensis (Castellani) Hau- duroy et al. (Bacillus negombensis Castellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 262; Hauduroy et al.. Diet. d. Bact. Path., 1937, 490.) Shigella oxy genes (Ford) Bergey et al. (Bacterium oxygenes Ford, Studies from the Royal Victoria Hospital, Montreal, 1, No. 5, 1903, 47; Eberthella oxygenes Bergey et al.. Manual, 1st ed., 1923, 228; Bergey et al.. Manual, 3rd ed., 1930, 360.) From feces. Shigella piscatora Bois and Roy. (Naturaliste Canadien, 71, 1945, 259.) From the intestine of a codfish (Gadus callarias L.). Shigella tangallensis (Castellani) Hau- duroy et al. (Bacillus tangallensis Cas- tellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 266; Hauduroy et al.. Diet. d. Bact. Path., 1937, 497.) From feces. Shigella tarda (Castellani) Hauduroy et al. (Bacillus tardus Castellani, 1917; see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 954; Hauduroy et al., Diet. d. Bact. Path., 1937, 497.) FAMILY_PARVOBACTERIACEAE 545 FMIILY XI. PARVOBACTERIACEAE RAHN.* (Cent. f. Bakt., II Abt., 96, 1937, 281.) Small, motile or non-motile rods. Gram-negative. Some will grow on ordinary media, but the majority either require or grow better on media containing body fluids or growth-promoting substances. Some invade living tissues. Usually do not liquefy gelatin. Xo visible gas formed in the fermention of carbohydrates. Infection in some cases may take place by penetration of organisms through mucous membranes or skin. Parasitic to pathogenic on w^arm-blooded animals, including man. Key to the tribes of family Parvobacteriaceae, I. Usually grow on ordinary media. A. Aerobic to facultative anaerobic. 1. Show bipolar staining. JMajority ferment carbohydrates. Tribe I. Pasteurelleae, p. 545. 2, Do not show bipolar staining. None ferment carbohydrates. Tribe II. Brueelleae, p. 560. B, Aaaerobic, Tribe III. Bacteroideae, p. 564. II. On first isolation dependent on some factor or factors contained in blood or plant tissues. Aerobic to anaerobic. Tribe IV. Hemophileae, p. 584. THIBE I. PASTEURELLEAE CASTELLANI AND CHALMERS. (Man. Trop. Med., 3rd ed., 1919, 943.) Small, motile or non-motile, ellipsoidal to elongated rods showing bipolar staining. Key to the genera of tribe Pasteurelleae. I. Milk not coagulated. A. Causes hemorrhagic septicemia, pseudotuberculosis, tularemia or plague. Genus I. Pasteur ella, p. 546. II. Milk coagulated slowly and sometimes digested. A. Causes glanders or glanders-like infections. Genus II. Malleomyces, p. 5.54. III. Milk unchanged to slightly acid. A. Associated with actinomycosis in cattle and in man. Genus III. Actinobacillus, p. 556. ''" Revised by Prof. E. G. D. ]\Iurray, ^IcGill University, Montreal, Canada with the collaboration of Prof. Karl F. Mej'er, Hooper Foundation, San Francisco, California; Prof. W. A. Hagan, Cornell University, Ithaca, New York; Dr. Alice C. Evans and Dr. Margaret Pittman, National Institute of Health, Washington, D. C; Prof. I. F. Huddleson, Michigan State College, East Lansing, Michigan; and others, December, 1938. 546 MANUAL .QF DETERMINATIVE fiAfiTERlOLOGY . . Genus I. Pasteurella Trevisan* {Octopsis Trevisan, Atti della Accad. Fisio-Medico-Statistica, Milario, Ser. 4, 3, 1885, 102; Trevisan, Rendiconti Reale Instituto Lombardo di Scienze e Lettere, 1887, 94; Coccobacillus Gamaleia, Cent. f. Bakt., 4, 1888, 167; Eucystia Enderlein, Sitzber. Gesell. Naturf. Freunde, Berlin, 1917, 317.) Named for Louis Pasteur, the French scientist. Small, Gram-negative, ellipsoidal to elongated rods showing bipolar staining by spe- cial methods; aerobic, facultative; may require low oxidation-reduction potential on primary isolation; majority ferment carbohydrates but produce only a small amount of acid; no or slight lactose fermentation; no gas production; gelatin not liquefied; milk not coagulated; parasitic on man, other mammals and birds. The type species is Pasteurella multocida (Lehmann and Neumann) Rosenbusch and Merchant. Key to the species of genus Pasteurella. I. Growth on ordinary media. Growth in milk. A. Non-motile and non-flagellated at 18° to 26°C. No change or slight acid in milk without coagulation. 1. Indole and H2S produced. No growth in bile. Sorbitol fermented. No hemolysis on blood agar. 1. Pasteurella multocida. 2. Indole not formed. Hemolysis produced on blood agar. 2. Pasteurella hemolytica. 3. Neither indole nor HoS produced. Growth in bile. Sorbitol not fermented. No hemolysis. 3. Pasteurella pestis. B. Motile and flagellated at 18° to 26°C. Milk alkaline. Hydrogen sulfide pro- duced. Indole not formed. 4. Pasteurella pseudotuberculosis. II. No growth on plain agar or in liquid medium without special enrichment. No growth in milk. 5. Pasteurella tularensis. 1. Pasteurella multocida (Lehmann dum Lehmann and Neumann, ibid., 196; and Neumann) Rosenbusch and Mer- Bacillus plurisepticus and Bacterium chant. (Virus der Wildseuche, Hueppe, avicidwn Kitt, in Kolleand Wassermann, Berlin, klin. Wochnschr., 23, 1886, 797; Handb. d. path. Mikroorg., 1 Aufl., 2, Bact^rie ovoide, Lignieres, Recueil de IQOZjb&l; Bacillus pleuriseplicus ^ovA^nv, Mdd. Vdt^r., 75, 1898, 836 (Bull. Soc. General Bact., 1st ed., 1908, 289; BaaWws Centr. Mdd. V^ter., N. S. 60, 1898, 836); bipolaris septicus Hutyra, in Kolle and Bacillus septicaemiae haemorrhagicae Wassermann, Handb. d. path. Mikroorg., Sternberg, Man. of Bact., 1893, 408; 2 Aufl., 6, 1913, 67; Bacillus bipolaris Bacterium septicaemiae hemorrhagicae plurisepticus Hutyra, ibid.; Pasteurella Lehmann and Neumann, Bakt. Diag., septica Topley and Wilson, Princip. 2 Aufl., 2, 1899, 194; Bacterium multoci- Bact. and Immun., 1st ed., 1, 1931, 488; * Rearranged by Mrs. Eleanore Heist Clise, New York State Experiment Station, Geneva, New York, in accordance with the suggestions of Mr. Philip C. Harvey and Dr. Mark Welsh, Pearl River, New York, November, 1945. FAMILY PARV0BACTP:EIACEAE 547 Pasteurella plurisepiica Gay et al., Agents of Disease and Host Resistance, 1935, 730; Rosenbusch and Merchant, Jour. Bact., 37, 1939, So.) From Latin, killing many. The following are regarded as identical 'dth the above but are arranged here iccording to source : Pasteurella bollingeri Trevisan. (Mi- croparasiten bei eine neue Wild- und Rinderseuche, Bollinger, tjber eine neue Wild- und Rinderseuche unsw., IVIiin- chen, 1878; Bacterium bipolare multo- ciduvi Kitt, Sitz. Gesell. Morphol. u. Physiol., Munchen, 1, 1885, 24; Trevisan, 1 generi e le specie delle Batteriacee, 1889, 21; Bacillus bovisepticus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 421 ; Bacterium boi'isepticus Chester, Ann. Rept. Del. Coi. Agr. Exp. Sta., 9, 1897, 81; Bacterium multocidum Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 196; La Pasteurella bo- vine, Lignieres, Recueil de M^d. Vet^r., 77, 1900, 537; Bacillus bipolaris bovisep- ticus Hutyra, in KoUe and Wassermann, Handb. d. path. Mikroorg., 2 Aufl., 6, 1913, 67; Pasteurella boviseptica Holland, Jour. Bact., 5, 1920, 224; Pasteurella bovium Hutyra, in KoUe, Kraus und Uhlenhuth, Handb. d. path. Mikroorg., 3 Aufl., 6, 1927-1929, 487; Pasteurella ferarum Hauduroy et al.. Diet. d. Bact. Path., 1937, 316.) From domestic cattle and deer. Pasteurella avicida (Gamaleia) Trevi- san. (Microbe du cholera des poules, Pasteur, Compt. rend. Acad. Sci., Paris, 90, 1880, 239, 952 and 1030 ; Granules of fowl cholera, Salmon and Th. Smith, U. S. Dept. Agr. Ann. Rept., 1880, 438; Micrococcus cholerae gallinarum Zopf, Die Spaltpilze, 3 Aufl., 1885, 57; Octopsis cholerae gallinarum Trevisan, Atti della Accad. Fisio-Medico-Statistica, Milano, Ser. 4, 3, 1885, 102; Bacillus cholerae gallinarum Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 253; Bacterium cholerae gallinarum Schroeter, Kryptogamen Flora von Schlesien, 3, 1, 1886, 155; Hiihnercholerabacterien, Kitt, Cent. f. Bakt., 1, 1887,305; Pasteurella cholerae gallinarum Trevisan, Rendiconti Reale Institute Lombardo di Scienze e Lettere, 1887, 94; Coccobacillus avicidus Gama- leia, Cent. f. Bakt., 4, 1888, 167; Trevisan, I generi e le specie delle Batteriacee, 1889, 21 ; Bacterium avicidum Kitt, ac- cording to Chester, Man. Determ. Bact., 1901, 135; Bacterium cholerae Chester, idem ; Bacillus avisepticus and Bacterium avisepticum Kitt, in Kolle and Wasser- mann, Handb. d. path. Mikroorg., 1 Aufl., 2, 1903, 544; not Bacillus avi- septicus Chester, loc. cit., 220; Pasteu- rella avium Kitt, loc. cit., 562; Pasteu- rella gallinae Besson, Practical Bac- teriology, London and New York, 1913, 447 ; Pasteurella cholerae-gallinarum Winslow et al., Jour. Bact., 2, 1917, 561; Bacillus cholerae-gallinarum Holland, Jour. Bact., 5, 1920, 217; Pasteurella aviseptica Holland, ibid., 224.) From fowls. Pasteurella cuniculicida (Fliigge) Trevisan. (Septiciimiebacterien, Gaff- ky, Mit. kaiserl. Gesundheitsamte, 1, 1881, 98; Bacillus cuniculicida Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 251; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 21; Bacterium septichae- miae Schroeter, Kryptogamen Flora von Schlesien, 3, 1 , 1889, 155 ; Bacterium cunic- ulicida Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 80; not Bacterium cuniculicida Chester, Man. Determ. Bact., 1901, 140; Bacillus cuniculisepticus Kitt, in Kolle and Wassermann, Handb. d. path. Mikroorg., 1 Aufl., 2, 1903, 562; Bacterium lepisepticum Ferry and Hos- kins. Jour. Lab. and Clin. Med., 5, 1920, 311; Bacillus bipolaris septicus and Bacillus lepisepticus Ford, Textb. of Bact., 1927, 591; Pasteurella lepiseptica Holland, Jour. Bact., 5, 1920, 221; Pas- teurella cuniculi Schiitze, Med. Res. Council, Syst. of Bact., London, 4, 1929, 469; Bacterium leporisepticum Haudu- roy et al., Diet. d. Bact. Path., 1937, 314.) From rabbits. Pasteurella suilla Trevisan. (Roth- laufstabchen, Loeffler, Arb. kaiserl. Ge- 548 MANUAL OF DETERMINATIVE BACTERIOLOGY sundheitsamte, 1, 1886, 51; Rothlauf- bacillen, Schiitz, ibid., 74; Bacillus of swine plague, Salmon, Rept. U. S. Dept. Agr., Bur. An. Ind., 1886, 87; Bacillus parvus ovatus Fltigge, Die Mikroorganis- men, 2 Aufl., 1886, 273; Trevisan, Reale Institute Lombardo d. Sci. e Let. Rend., Ser. 2, 20, 1887, 94; Bacterium suicida Migula, in Engler and Prantl, Natiirl. Pflanzenfam., 1, la, 1895, 27; Bacillus suisepticus Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., !i, 1896, 419; Bac- terium suisepticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 80; La Pasteurella porcine, Lignieres, Recueil de Mdd. Vdt6r., 77, 1900, 391 ; Pasteurella suiseptica Holland, Jour. Bact., 5, 1920, 220; Pasteurella suum Hutyra, in Kolle, Kraus and Uhlenhuth, Handb. d. path. Mikroorg.,-3 Aufl., 6, 1927-1929, 487.) From swine. Bacterium bovicida Migula. (Microbo del barbone dei bufali, Oreste Armanni, Atti. d. R. Istit. d'incoragg. alle scienze natur. ecenom. e technol., 1887; Letta nella tornata Accad., Sept. 16, 1886; Cent. f. Bakt., 2, 1887, 50; Atti della Commissione per le malattie degli ani- mali, 121, 1887; Migula, Syst. d. Bakt., 2, 1900, 366; Pasteurella bubalseptica Kelser, Man. Vet. Bact., 1st ed., 1927, 195; Bacillus bubalsepticus Kelser, ibid.; Bacillus bipolaris bubalisepticus Haudu- roy etal.. Diet. d. Bact. Path., 1937, 312.) From buffaloes. Pasteurella vituliseptica (Kitt) Ford. (Bacillus vitulisepticus Kitt, in Kolle and Wassermann, Handb. d. path. Mi- kroorg., 1 Aufl., 2, 1903, 562; Bacterium vitulisepticum Lehmann and Neumann, Bakt. Diag., 5 Aufl., 2, 1912, 282; Ford, Textb. of Bact., 1927, .597.) From calves. Pasteurella muricida Meyer and Batch- elder. (Meyer and Batchelder, Jour. Inf. Dis., 39, 1926, 386; Pasteurella muriseptica Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 482; not Pasteurella muriseptica Bergey et al., 1st ed., 1923, 265 (Bacillus murisepticus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 250; Erysipelothrix muriseptica Bergey et al., Manual, 2nd ed., 1925, .380).) From wild rats. Bacillus bipolaris der nialignen Meer- schwcinchen-Phlegmasie of Heymannand Kyriasides, Ztschr. f. Hyg., lU, 1932, 119 (Klebsiella caviae Hauduroy et al.. Diet, d. Bact. Path., 1937, 261) is stated by the original authors to be closely related to this organism. Plasaj and Pribram (Cent. f. Bakt., I Abt., Orig., 87, 1921, 1) also present a classification of the hemorrhagic septi- cemia bacteria. Description from Schiitze (Med. Res. Council, Syst. of Bact., London, 4, 1929, 451 ) who prepared it from studies of 230 strains described by 17 authors during the years 1908-1926. Short ellipsoidal rods: 0.3 to 1.25 microns in length, occurring singly, in pairs, rarely in chains. Show bipolar staining. Non -motile. Gram-negative. Gelatin: No liquefaction. Agar: Fine translucent growth. Char- acteristic odor. Broth: Uniform turbidity. Charac- teristic odor. Milk: No change in reaction. No coagulation. Potato : No visible growth . Indole is formed. Nitrites are produced from nitrates. Hydrogen sulfide is produced. No hemolysis on blood agar. Acid but no gas from glucose, mannitol (usually), sucrose, fructose, sorbitol, galactose, mannose, xylose (usually) and trehalose (usually). No acid from lac- tose, dulcitol, arabinose (usually), amyg- dalin, maltose (usually), raffinose, rhamnose, adonitol, dextrin, inulin, glyc- erol, salicin (usually) or erythritol. Optimum temperature 37°C. Killed at temperatures above 45°C. Aerobe to facultative anaerobe. Three serological types have been found on the basis of agglutination tests (Little and Lyon, Amer. Jour. Vet. Res., 4, 1943, 110). FAMILY PARVOBACTERIACEAE 549 Virulent for laboratory animals, espe- cially mice and rabbits. Distinctive characters: Grows on or- dinary media. Bile salts inhibit growth. Source : From numerous domestic ani- mals and fowls, including cat, dog, cattle, horse, goat, sheep, pig, rabbit, chicken, and from reindeer, buffalo, rat, etc. Habitat : The cause of hemorrhagic septicemia in birds and mammals. 2. Pasteurella hemolytica Newsomand Cross. (Jour. Amer. Vet. INIed. Assoc, 80 (N.S. 33), 1932, 715.) From M. L., hemolytic. Bipolar staining. Blood agar: Hemolysis. Indole not formed. Acid from dextrin, fructose, galactose, glucose, glycerol (usualljO, inositol, lac- tose (usually), maltose, mannitol, raf- finose, sorbitol, sucrose and xylose. Xo acid from arabinose, dulcitol, inulin, mannose, rhamnose or salicin. Xo cross-agglutination between Pas- teurella multocida and this species. Avirulent for rabbits. Source : Twenty strains isolated from pneumonia in sheep and cattle. Habitat : Occurs in pneumonia of sheep and cattle. .3. Pasteurella pestis (Lehmann and X'eumann) Holland. (Bacille de la peste, Yersin, Ann. Inst. Past., 8, 1894, 666; Pest Bacillus, Aoyama, Ztschr. f. Hyg., 21, 1895, 165; Bacterium pestis Lehmann and Xeumann, Bakt. Diag., 1 Aufl., 2, 1896, 194; Bacillus pestis biibonicae Kruse, in Fliigge, Die INIikroorganismen, 3 Aufl., 2, 1896, 429; Bacterium pestis bubonicae Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 81; Bacillus pestis Migula, Syst. d. Bakt., 2, 1900, 749; Eucystia pestis Enderlein, Sitzber. Gesell. Xaturf. Freunde, Berlin, 1917, 317; Holland, Jour. Bact., 5, 1920, 219; Coccobacillus yersini Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 20.) From Latin pestis, plague. Rods: 1.0 by 2.0 microns, occurring singly. X'on-motile. Polar staining. Characteristic bladder, safetj^-pin and ring involution forms. Gram-negative. Gelatin colonies: Flat, gray, with granular margin. Gelatin stab: Flat surface growth. Arborescent growth in stab. Xo lique- faction. Agar colonies: Grayish-white, translu- cent, irridescent, undulate. Agar slant : Growth grayish, viscid, thin, moist, translucent. Growth slow, favored by the addition of blood or so- dium sulfite. Broth : Turbid or clear with fiocculi in the fluid. Old cultures show a pellicle with streamers into the fluid (stalac- tites). Becomes alkaline more slowly than Pasteurella pseudotuberculosis. See Bessonowa and Lenskaja, Cent. f. Bakt., I Abt., Orig., 119, 1930,430. Litmus milk: Slightly acid or un- changed. Xo coagulation. Potato: Scanty, grayish growth. Indole not formed. Lactose and rhamnose not attacked. Variable action on glycerol. Xitrites are produced from nitrates. Temperature relations: Optimum 25° to 30°C. Minimum 0°C. Maximum 43° to 45 °C. Aerobic, facultative. Source : Buboes, blood, pleural effusion, spleen and liver of infected rodents and man. Sputum in pneumonic plague. Infected fleas. Habitat : The causative organism of plague in man, rats, ground squirrels and other rodents. Infectious for mice, guinea pigs and rabbits. Transmitted from rat to rat and from rat to man by the infected rat flea. Xote: Pasteurella pestis and Pas- teurella pseudotuberculosis are not defi- nitely distinguishable by serological methods (Schiitze, Med. Res. Council, Syst. of Bact., London, 4, 1929, 478, and Wu Lien-teh, in Chun, Pollitzer and Wu, "Plague," Xational Quarantine Service, Shanghai, 1936). Malachite-green broth slowly decolorized by Pasteurella pestis 550 MANUAL OF DETERMIXATIVE BACTERIOLOGY and quickly by P. pseudotuberculosis; same for methylene blue, Janus green and thionin. No growth on Bessonowa media (pH 5.9). See Yersinia, p. 703. 4. Pasteurella pseudotuberculosis (Eisenberg) Topley and Wilson. (Ba- cillus der pseudotuberculose, Ffeiffer, Ueber die bacillare Pseudotuberculose bei Nagetieren, Leipzig, 1889 ; Bacillus pseu- dotuberculosis Eisenberg, Bakt. Diag., 3 Aufl., 1891, 294; Streptobacillus pseudo- tuberculosis rodentium Preisz, Ann. Inst. Past., 8, 1894, 231; Bacterium pseudo- tuberculosis rodentium Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 362; Bacillus pseudotuberculosis roden- tium Lehmann and Neumann, ibid., 429; Bacterium pseudotuberculosis Migula, Syst. d. Bakt., 2, 1900, 374; Corynebac- terium rodentium Bergey et al.. Manual, 1st ed., 1923, 386; Corynebacterium pseudotuberculosis Bergey et al., Manual, 2nd ed., 1925, 394; Topley and Wilson, Princip. Bact. and Immun., 1st ed., 2, 1931, 825; Corynebacterium pseudotu- berculosis rodentium Kelser, IVIan. Vet. Bact., 2nd ed., 1933, 319; Malleomyces pseudotuberculosis rodentium Pribram, Klassifikation d. Schizomyceten, 1933, 93.) Latinized, false tuberculosis. Bacillus tuberculosis zoogloeicae Malas- sez and Vignal (Ann. de Physiol., 1883, 370) is considered identical with this species by Hauduroy et al. (Diet. d. Bact. Path., 1937, 162). The original tabular description by Eisenberg, Bakt. Diag., 3 Aufl., 1891, 294 is very incomplete. Description taken from Topley and Wilson, Princip. Bact. and Immun., 2nd ed., 1936, 607 and Bessonowa, Lenskaja and Molodtzowa, Office Internat. d'Hyg. Publ., 29, 1937, 2106. Small rods : Variable in size and shape. Ellipsoidal or coccoid forms 0.8 by 0.8 to 2.0 microns, with rounded ends, occur- ring singly. Rod-shaped forms 0.6 by 1.5 to 5.0 microns, with rounded ends, occurring singly, in groups or in short chains. Occasionally long curved fila- ments. Motile (Weitzenberg, Cent. f. Bakt., I Abt., Orig., 133, 1935, 343). Non-acid-fast. Gram-negative. Gelatin stab : After 7 days at 22°C, good filiform growth extending to bottom of tube. No liquefaction. Agar colonies: After 24 hours at 37°C, circular, 0.5 to 1.0 mm in diameter, umbonate, granular, translucent, gray- ish-yellow, butyrous; edge entire; dull, finely granular or beaten-copper surface ; differentiated into a raised, more opaque center and a flat, clearer periphery with radial striation. Agar slant: After 48 hours at 37°C, growth moderate, confluent, raised, gray- ish-yellow, translucent, with glistening, wavy or boaten-copper surface and an irregularl}^ lobate edge. Blood agar plate : Good growth. No hemolysis. Broth : After 24 hours at 37°C, mod- erate growth with moderate turbidity which later clears. Viscous sediment. Incomplete surface and ring growth. Becomes alkaline more rapidly than Pasteurella pestis. Potato : After 7 days at 22°C, a thin yellowish membrane which later turns brown. Indole not formed. Litmus milk : Usually slightly alkaline. Nitrites produced from nitrates. Ammonia is produced. Acid but no gas from glucose, maltose, mannitol, salicin, arabinose, xylose, rhamnose and glj^cerol. Sometimes acid from sucrose. Hydrogen sulfide produced. Catalase positive. Methyl red positive. Methylene blue is reduced. Voges-Proskauer test negative. Temperature relations : Optimum 30°C. Minimum 5°C. Maximum 43 °C. Ther- mal death point 60°C for ten minutes. Pathogenicity : The cause of sponta- neous disease in rabbits, rats and guinea pigs. Infectious for mice, rats, dogs, cats and horses. Aerobic, facultative. FAMILY PARVOBACTERIACEAE 551 Source : From a guinea pig inoculated with material from a horse suspected of having glanders. Habitat : Lesions in natural disease in animals. Causes pseudotuberculosis in rodents, especial!}' guinea pigs. 5. Pasteurella tularensis (McCoy and Chapin) Bergey et al. {Bacterium tula- rense McCoy and Chapin, Jour. Inf. Dis., 10, 1912, 61 ; McCoy and Chapin, Public Health Bull. 53, U. S. Treas. Dept., Public Health Service, 1912, 17; Bacillus tularense Vail, The Ophthalmic Record, 23, 1914, 487; also Francis, U. S. Hygienic Lab. Bull. 130, 1922; Bergey et al., Manual, 1st ed., 1923, 267; Bru- cella tularensis Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 509; Coccohacterium tularense Galli- Valerio, Schweiz. med. Wochnschr., 68, 1938, 1206.) From Tulare, the county in California in which the disease was first observed. Description taken from McCoy and Chapin {loc. cit.) and Francis (loc. cit.). Further revision by Francis, 1947. Equal numbers of cocci and rods; 0.2 by 0.2 to 0.7 micron, occurring singly. Bipolar staining may occur. Capsules rare or absent. Extremely pleomorphic (Hesselbrock and Foshay, J. Bact. 49, 1945, 209) Non-motile. Gram-negative. Xo growth on plain agar or in liquid media without special enrichment. (Tamura and Gibby, J. Bact. 45, 1943,361) Filterable through Berkefeld filters. Growth occurs on coagulated egg-yolk (McCoy and Chapin, loc. cit.), on blood- glucose -cystine agar (Francis, loc. cit.), on blood agar, glucose-blood agar and glucose serum agar. The addition of fresh sterile rabbit spleen to the surface of the last three media favors the growth of the organism. Forms minute viscous colonies after 2 to 5 days which may attain a diameter of 4 mm if well separated. Growth readily emulsifiable. Growth on blood media is gray. May cause green discoloration of the blood. Rough, smooth and mucoid variants liave not been reported. Slight acid without gas may be pro- duced from glucose, glycerol, maltose, mannose, fructose and dextrin. Growth .soluble in sodium ricinoleate. Hj^drogen sulfide produced in a cystine medium. Aerobic. No growth anaerobically. Optimum temperature 37 °C. Thermal death point 56°C for ten minutes. Sur- vives best at low temperatures, even -70°C. Pathogenicity : Penetrates unl^roken skin to cause infection. Buboes and areas of necrosis produced in human and animal tissue. Infectious for man and most rodents, including rabbits, guinea pigs, rats, mice, squirrels, ground hogs, muskrats, beavers, water rats and lemmings. Source: Originally isolated from Cali- fornia ground squirrels and later from more than 30 other forms of wild life in the United States and elsewhere. Found in lesions in man and animals with natu- ral or experimental infections. E^spe- cially the liver, blood, lymph nodes, and spleen of animals. Habitat : The cause of tularemia in man and transmitted from wild animals to man by blood-sucking insects, by con- tact with infected animals, or bj' drinking water. Disease known in North America, Japan, Russia, Norway, Sweden, Austria, Turkey, Czechoslovakia and Central Germany. See Burroughs, Holdenreid, Longanecker and Meyer, Jour. Inf. Dis., 76, 1945, 115 for a complete list of known vertebrate hosts. Appendix: The following organisms may be identical with some of those listed above or related to them: Bacillus coscoroba Tretrop. (Tretrop, .\nn. Inst. Past., U, 1900, 224; not Bacil- lus coscoroba MacConkey, Jour. Hyg., 6, 1906, 397.) The cause of swan cholera in the Antwerp Zoological Garden. Tretrop 's description is that of a Pas- teurella as is pointed out by Castellani 552 MANUAL OF DETERMINATI\T: BACTERIOLOGY and Chalmers (Man. Trop. Med., 3rd ed., 1919, 941). The organism described by Tr^trop clearly was not the same as that in the culture sent by Binot of the Pas- teur Institute to MacConkey and de- scribed by him {loc. cit.) as a member of the coliform group. Because of Mac- C!onkey's studies, the Binot culture has been accepted as determining the nature of Bacillus coscoroba in many subsequent studies of the coliform group, e. g., Bergey and Deehan, Jour. Med. Res., 19, 1908, 182; Levine, Amer. Jour. Pub. Health, 7, 1917, 785; Winslow, Kligler and Roth- berg, Jour. Bact., 4, 1919, 485; Bergey et al.. Manual, 1st ed., 1923, 204; etc. Bacillus cuniculicida Migula. (Bacil- lus der Kaninchenseptikamie, Eberth and Mandry, Arch. f. path. Anat., 121, 1890; Bacillus cuniculicida mobilis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 406; Bacterium cuniculicida mobilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 132; Migula, Syst. d. Bakt., 2, 1900, 757; not Bacillus cuniculicida Fliigge, Die Mikroorganis- men, 2 Aufl., 1886, 251.) From perito- neal exudate of a rabbit. Bacillus muslelaecida Trevisan. (Ba- cillus der Frettchenseuche, Eberth and Schimmelbusch, Fortschr. d. Med., 6, 1888, 295; also see Arch. f. path. Anat., 115, 1889, 282; Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Pas- teurella muslelaecida DeToni and Trevi- san, in Saccardo, Sylloge Fungorum, 8, 1889, 996; Bacillus micstelae septicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl.,^, 18m, 405; Bact eriu7n mustelae septicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 138; Bacillus mustelae Migula, Syst. d. Bakt., 2, 1900, V and 756.) From a disease of ferrets. Bacterium, anatis Migula. (Bact^ries du cholera des canards. Cor nil and Tou- pet, Compt. rend. Acad. Sci. Paris, 106, 1888, 1747; Bacillus cholerae anatum Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896,417; Migula, Syst. d. Bakt., 2, 1900, 364.) Regarded as the cause of duck cholera and very similar to, if not identical with, Pasleurella avicida. See Rettger and Scoville, Jour. Inf. Dis., 26, 1920, 220. From the blood and other organs of infected ducks. Bacterium cunicuU Migula. (Bacillus der Brustseuche des Ivaninchens, Beck, Ztschr. f. Hyg., 15, 1893, 363; Bacillus cunicidi pneumonicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 418; Bacterium cuniculi pneumonicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 84; Migula, Syst. d. Bakt., 2, 1900, 370; Bacterium beckii Chester, Man. Determ. Bact., 1901, 142.) Asso- ciated with a lung plague of rabbits. Bacterium haemorrhagicum (Kruse) Lehmann and Neumann. (Kolb, Arb. kaiserl. Gesundheitsamte, 7, 1892, 60; Bacillus haemorrhagicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 424; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 194.) From the mucous membranes of fever patients. Bacterium palumbarium Migula. (La bacterie de la maladie des palombes, Leclainche, Ann. Inst. Past., 8, 1894, 493; Bacillus cholerae columbarum Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 417; Bacterium cholerae colum- barum Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 84; Migula, Syst. d. Bakt., 2, 1900, 368; Bacterium colum- barum Chester, Man. Determ. Bact., 1901, 141.) Associated with an epidemic in wild pigeons. Bacterium phasianicida Klein. (Klein, Cent. f. Bakt., I Abt., Orig., 31, 1902, 76; Bacterium phasianidarum mo- bile Enders, Berl. tierarztl. Wchnschr., No. 23, 1902; abst. in Cent. f. Bakt., I Abt., Ref., 34, 1904, 384.) From an epidemic in pheasants (England). Had- ley, Elkins and Caldwell (Rhode Island Agr. Exp. Sta., Bull. 174, 1918, 28) state that this species (which they call B. phasianicida) belongs in the group of paratyphoids (Salmonella) . Bacterium purpurum Chester. (Ba- cillus of purpura-haemorrhagica, Babes, Septiche Proz. Kindesalters, Leipzig, 1889; Bacillus haemorrhagicus septicus FAMILY PARVOBACTERIACEAE 553 Kruse, in Fliigge, Die Mikroorganismeu, 3 Aufl., 2, 1896, 424; Bacterium haemor- rhagicus septicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Chester, Man. Determ. Bact., 1901, 143.) From a case of septicemia in man. Bacterium lizzonii Migula. (Bacillus der haemorrhagischen Infektion, Tizzoni and Giovannini, in Zeigler, Beitrage, 7, 1889, 300; Bacillus haemorrhagicus vele- nosus Kruse, in Fliigge, Die Milcroor- ganismen, 3 Aufl., 2, 1896, 425; Bac- terium haemorrhagicus velenosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Migula, Syst. d. Bakt., 2, 1900, 386; Bacterium velenosum Chester, Man. Determ. Bact., 1901, 144.) From the blood of a child having a hemorrhagic infection. Bacterium vassalei Migula. (Tizzoni and Giovannini, in Zeigler, Beitrage, 7, 1889 ; Bacillus haemorrhagicus nephritidis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 424; Bacterium haemor- rhagicus nephritidis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Migula, Syst. d. Bakt., 2, 1900, 387; Bacterium nephritidis Chester, Man. Determ. Bact., 1901, 145.) Isolated by Vassale from a case of hemorrhagic nephritis. Pasieitrcllu houffardi Commes. (Commes, 1919; quoted from Xeveu- Lemaire, Precis de Parasitol. Hum., 5th ed., 1921, 21.) The cause of a human pasteurellosis observed bj* Bouffard at Bamako in 1909. Pasteurella caniseptica Hauduroy etal. (Pasteurella du chien, Lignieres, Recueil de Med. Vet., 77, 1900, 469; Bacterium canicida Lehmann and Xeumann, Bakt. Diag., 4 Aufl., 2, 1907, 277; Hauduroy et al.. Diet. d. Bact. Path., 1937, 312.) From dogs. Pasteurella capriseptica (Lanfranchi and Pacchioni) Hauduroy et al. {Bacillus pneumoniae caprae Nicolle and Refik- Bey, Ann. Inst. Past., 10, 1896, 321; Pasteurella du chevre, Lignieres, Recueil de M(5d. V^t^r., 77, 1900, 536; Bacillus caprisepticus Lanfranchi and Pacchioni ^ 1926; Bacillus bipolaris caprisepticus Chefik Kolayi and Raif , 1935 ; Hauduroy et al.. Diet. d. Bact. Path., 1937, 313.) From hemorrhagic septicemia in goats. Pasteurella caviae Hauduroy et al. (Gate and Billa, Compt. rend. Soc. Biol., Paris, 99, 1928, 814; Hauduroy et al.. Diet. d. Bact. Path., 1937, 313.) From a guinea pig with a tuberculosis-like disease. Pasteurella caviseplica (Sehwer) Hau- duroy et al. (Pasteurella du cobaye, Phisalix, Compt. rend. Soc. Biol., Paris, /, 1898, 761; Bacterium cavisepticum Sehwer, Cent. f. Bakt., I Abt., Orig., 33, 1902, 47; Hauduroy et al., Diet. d. Bact. Path., 1937, 314.) From hemorrhagic septicemia in guinea pigs. Pasteurella desmodilli Pirie. (Pub. So. African Inst. Med. Res., 4, 1929, 191.) Pasteurella equiseptica Kelser. (Ba- eille de la septicemic hemorragique du cheval, Lignieres, Bull. Soc. Centr. d. Med. Vet., 15, 1897, 437 and 16, 1898, 849; La Pasteurella equine, Lignieres, Recueil de Med. V^t., 77, 1900, 524; Kelser, Man. Vet. Bact., 1st ed., 1927, 191; Bacillus cquisepticus Kelser, ibid. ; Bacillus pneu- moniae equi Poels.) From horses. Pasteurella felis (Migula) Hauduroy et al. (Bacillus salivarius septicus felis Fioeca, Ann. d. Instit. d'igiene d. Univ. di Roma, 2, 1892 and Cent. f. Bakt., 11, 1892, 406; Bacillus felis septicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 423; Bacterium felis septi- cus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 81; Pasteurella du chat, Lignieres, Recueil de Med. Vet., 77, 1900, 493; Bacterium felis Migula, Syst. d. Bakt., 2, 1900, 375; Hauduroy et al.. Diet. d. Bact. Path., 1937, 316.) From the sputum of a cat. Pasteurella mastidis (Miessner and Schoop) Hauduroy et al. (Stiibchenbak- terium, Dammann and Freese, Deut. tierarztl. Wchnschr., 15, 1907, 165; Bipolar organism of the Pasteurella group, Leyshon, Vet. Jour., 85, 1929, 299; Bacterium mastitidis Miessner and Schoop, Deut. tierarztl. Wchnschr., 40, 554 MANUAL OF DETERMINATIVE BACTERIOLOGY 1932, 69; Pasteurella, Marsh, Jour. Amer. Vet. Med. Assoc, 81 (N.S. 34), 1932, 376; Bacterium ovinum Haupt, Cent. f. Bakt., I Abt., Orig., 123, 1932, 365; Hauduroy et al.. Diet. d. Bact. Path., 1937, 316.) The cause of infec- tious mastitis of ewes. Pasteurella necrophora Hauduroy et al. (Bacille de la necrose infectieuse des Canaris, Cornell, The Vet. Record, 84, 1928, 350; Hauduroy et al., Diet. d. Bact. Path., 1937, 318.) From domestic canaries. Pasteurella oviseptica Hauduroy et al (Galtier, Jour. d. m^d. v6t. et d. zoot., 1889-1890, 58, 113 and 481; La Pasteurella ovine, Lignieres, Recueil de M^d. Y6t4iv., 77, 1900, 529; Bacillus bipolaris ovisepticus Hutyra, in Kolle and Wassermann, Hand. d. path. Mikro- org-., 2 Aufl., 6', 1913, 67; Hauduroy et al., Diet. d. Bact. Path., 1937, 319.) From sheep. Pasteurella pericarditis Hauduroy et al. {Bacterium cavarum pericarditis Roth, Acta Pathol, et Microb. Scand., 11, 1934, 335; Hauduroy et al., Diet, d. Bact. Path., 1937, 319.) From guinea pigs. Pasteurella strasburgensis Hauduroy et al. (Coccobacille de Strasbourg, Debre, Compt. rend. Soc. Biol., Paris, 82, 1919, 224; Hauduroy et al., Diet, d. Bact. Path., 1937, 323.) From a case of purulent pleurisy. Pfeifferella anatipestifer Hendrickson and Hilbert. (Hendrickson and Hilbert, The Cornell Veterinarian, 22, 1932, 239; Hemophilus anatipestifer Hauduroy et al.. Diet. d. Bact. Path., 1937, 247.) From a septicemic disease of ducks. Genus II. Malleomyces Pribram.* {Cladascus Enderlein (in part), Sitzber. Gesell. Naturf. Freunde, Berlin, 1917,316; Pfeifferella Buchanan, Jour. Bact., 3, 1918, 54; Pribram, Klassification der Schizo- myceten, Leipzig, 1933, 11 and 93; Loefflerella Gay et al., Agents of Disease and Host Resistance, Indianapolis, 1935, 782.) From Latin malleus, glanders and myces, fungus. Because Pfeifferella was proposed inadvertently (Buchanan, Gen. Syst. Bact., 1925, 420) and because of a general feeling that it is inappropriate, Malleomyces Pribram is used as the earliest suitable name for this genus. The indefinite description of an organism {Malleomyces equestris) by Hallier (Ztschr. f. Parasitenkunde, 1870, 119) as the cause of glanders has not previously caused confusion and need not do so in the future. Short rods, with rounded ends, sometimes forming threads and showing a tendency toward branching. Motile or non-motile. Gram-negative. Tendency to bipolar staining. Milk slowly coagulated. Gelatin may be liquefied. Specialized for para- sitic life. Grow well on blood serum and other body fluid media. The type species is Malleomyces mallei (Flligge) Pribram. Key to the species of genus Malleomyces. I. Carbohydrates not fermented. Honey-like colonies on potato. Glycerol agar colonies slimy or tenacious, translucent. N(jn-motile. 1. Malleomyces mallei. II. Carbohydrates fermented. Profuse, creamy growth on potato. Glycerol agar colonies iridescent, becoming corrugated. Motile. 2. Malleomyces pseudomallei. * Revised by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, December, 1938; further revision, December, 1945. FAMILY PARVOBACTERIACEAE OOO 1. Malleomyces mallei (Zopf) Pri- bram. (Rotzpilz, LofRer and Schlitz, Deutsche med. Wchnschr., Xo. 52, 1882; Bacillus mallei Zopf, Die Spaltpilze, 3 Aufl., 1885, 89; Rotzbacillus, Loffler, Arb. kaiserl. Gesundheitsamte, 1, 1886, 222; Bacterium mallei 'Sligula, in Engler and Prantl, Die natlirl. Pflanzenfam., /, la, 1895, 21; Corynehacterium mallei Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 366; Mycobacterium mal- lei Chester, Man. Determ. Bact., 1901, 353; Cladascus mallei Enderlein, Sitzber. Gesell. Xaturf. Freunde Berlin, 1917, 395; Pfeifferella mallei Buchanan, Jour. Bact., 3, 1918, 54; Sclerothrix mallei Vuillemin, Encyclopedic Mycolog., Paris, 2, Champignons Parasites, 1931, 135; Brucella mallei Pacheco, Revista da Sociedado paulista de IMedicina veteri- naria, 3, 1933, 1; Actinobacillus mallei Thompson, Jour. Bact., £6, 1933, 226; also Jour. Bact., 25, 1933, 44; Pribram, Klassification der Schizomyceten. Leip- zig and Vienna, 1933, 93; Loefflerella mallei Gay et al., Agents of Disease and Host Resistance, Indianapolis, 1935, 782.) From Latin malleus, glanders, a disease of horses. Bacillus ozenae Trevisan (Corr. Ser., 1884, n. 222) is identical with this species according to Trevisan (I generi e le specie delle Batteriacee, 1889, 13). Description largely from Kelser, INIan. Vet. Bact., 2nd ed., 1933, 325. Slender rods: 0.5 to 1.0 by 2.0 to 5.0 microns, with rounded ends, usually occurring singly, in pairs and in groups, but may grow into filaments. Branch- ing involution forms on glycerol agar. Show irregular staining. Bipolar stain- ing common. Non-motile. Gram-nega- tive . Gelatin: Poor growth. Usually' no liquefaction. May be slowly liquefied (Jordan, General Bact., 11th ed., 1935, 491). Agar colonies: Moist, grayish -white laj'er, translucent, ropy, with regular borders. Later become yellowish or yellowish-brown. Agar slants; Glistening, moist, ropy, grayish-white growth. Loffler's serum: Good growth. Moist, viscid, yellowish colonies develop after 36 to 48 hours. Broth: Turbid, sometimes with thin pellicle. Slimy or ropy sediment. Litmus milk: Coagulation usually occurs after a week with some acid pro- duction. Litmus maj' or may not be reduced. Potato : After 36 to 48 hours, pale yellow, hcney-drop-like colonies. Later becoming darker, reddish-yellow or choc- olate color. The medium sometimes has a faint greenish tinge around the growth. Indole not formed. Nitrites not produced from nitrates. Carbohydrates usuallj' not fermented. Some strains produce small amounts of acid from glucose. Optimum temperature 37°C. No growth below 20°C or above 44°C. Aerobic, facultative anaerobic. Common name : Glanders bacillus. Distinctive characters: Culture media of slightly acid reaction best suited for growth; addition of glycerol favors growth ; honej'-like growth on potato. Source : Isolated by Loffler and Schiitz from the liver and spleen of a horse. Lesions in animals and man. Habitat : The cause of glanders, affect- ing horses, man, sheep and goats. Trans- missible to dogs, cats, rabbits and guinea pigs. 2. Malleomyces pseudomallei (Whit- more) Breed. (Bacillus pseudomallei Whitmore, Jour. Hyg., 13, 1913, 1 ; Bacil- lus whitmori Stanton and Fletcher, Trans. 4th Cong. Far East Assn. Trop. Med., 2, 1921, 196; also Jour. Hyg., 23, 1925, 347; Pfeifferella pseudomallei Ford, Textb. of Bact., 1927, 294 ; Flavobacterium pseudomallei Bergey et al.. Manual, 3rd ed., 1930, 146; Sclerothrix whilmori Vuillemin, Encja^lop^die Mycolog., 2, Champignons Parasites, 1931, 136; -4c- finobacillus pseudom,allei Thompson, Jour. Bad., 26, 1933, 226; also Jour. Bact., 556 MANUAL OF DETERMINATIVIO BACTERIOLOGY 35, 1933, 44; LoeJJlerella whitmori Gay et al., Agents of Disease and Host Re- sistance, Indianapolis, 1935, 791 ; Breed, in Manual, 5th cd., 1939, 300.) Latin- ized, false glanders. Short rods : With rounded ends, occur- ring singly and in short chains, showing bipolar staining. Motile. Gram-nega- tive. Gelatin stab : Moderate, crateriform liquefaction. Agar colonies : Circular, slightly raised, thick, opaque, cream-colored with ir- regular margin. Glycerol agar slant : Wrinkled, thick, rugose, cream-colored growth. Broth: Turbid with pellicle. Litmus milk: Curdling with slowly developed acidity, pink sediment ; may be digested. Potato: Vigorous, cream-colored growth. Indole not formed. Acid from glucose, maltose, lactose, sucrose and mannitol. Blood serum slowly liquefied. Aerobic, facultative. Optimum temperature 37°C. Source : Lesions and blood in rats, guinea pigs, rabbits and man. Once from a transient nasal discharge in a horse (Stanton, Fletcher and Symonds) and once from a splenic abscess in a cow (Nicholls). Habitat: Glanders-like infection (meli- oidosis) in rats, guinea pigs, rabbits and in man in India, Federated Malay States and Indo-China. Appendix: The following may belong in this genus : Flavobacierium orchitidis Sherwood, Irwin and Marts. (Sherwood, Irwin (not Edwin) and Marts, Amer. Jour. Diseases of Children, 45, 1933, 446; Sherwood, Jour. Kansas Med. Soc, 34, 1933, 220.) From a case of meningitis. Sherwood (personal communication, 1945) now considers this organism iden- tical with Bacillus whitmori (MaUeo- myccs pseudomallei). See Manual, 5th cd., 1939, 53S for a description of this species. Malleomyces agliaceus Pribram. (Bacillo opale agliaceo, Vincenzi, Giorn. d. R. Accad. d. Med. Torino, 1890, No. 6; Pribram, Klassification der Schizomy- ceten, Leipzig, 1933, 93.) Cause of ))seudotuberculosis in frogs. Genus III. Actinobacillus Brumpt* (Precis de Parasitologic, Paris, 1st ed., 1910, 849.) Medium-sized, aerobic, Gram-negative rods which frequently show much pleomor- phism. Coccus-like forms frequent. Tendency to bipolar staining. Acid but no gas produced from carbohydrates. Grow best, especially when freshly isolated, under increased CO2 tension. Pathogenic for animals; some species attack man. The out- standing characteristic of the group is the tendency to form aggregates in tissues or culture which resemble the so-called sulfur granules of actinomycosis. The type species is Actinobacillus lignieresi Brumpt. 1. Actinobacillus lignieresi Brumpt. (Actinobacilo, Lignieres and Spitz, Bole- tin d. Agri. y Ganaderia, Buenos Aires, 11, 1902, 169; Actinobacillus, Lignieres and Spitz, Cent. f. Bakt., I Abt., Orig., 35, 1903, 294; Brumpt, Precis de Para- sitol., Paris, 1st ed., 1910, 849; Bacillus lignieri (sic) Mace, Traite de Bact^riolo- * Revised by Prof. W. A. Hagan, New York State Veterinary College, Ithaca, New York, December, 1938; further revision, December, 1945. FAMILY PARVOBACTERIACEAE 557 gie, 6th ed., 2, 1913, 743; Nocardia lig- nieresi Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 242; Discomyces lignieresi Brumpt, Pre- cis de Parasitol., Paris, 3rd ed., 1922, 993; Pasteurella lignieresi Vuillemin, Encj^clopedie Mycologique, Paris, 2, Champignons Parasites, 1931, 136.) Named for Lignieres, who first worked with this organism. Bacterium purifaciens Christiansen (JVIaanedsskr. f. Drylaeger, 29, 1917, 449; Pasteurella purifaciens Hauduroy et al., Diet. d. Bact. Path., 1937, 322) is regarded as identical with this organism by Tunnicliff (Jour. Inf. Dis., 69, 1941, 52). Rods: 0.4 bj'^ 1.0 to 15.0 microns. Cocco-baciDary forms frequent. Non- motile. Gram-negative. Gelatin: Growth sparse or fails. No liquefaction. Agar: Primary cultures usually suc- ceed best when the inoculum is intro- duced by stab. Serum agar is more favorable than plain. Surface colonies are small, bluish, translucent at first, later becoming opaque. Broth : Serum favors growth. Freshly isolated strains usually grow in form of small granules which adhere to sides of tube, leaving broth fairly clear. Later most strains grow diffusely, often form- ing a fragile pellicle. Litmus milk: Most strains cause no change. Sometimes slight acid. No co- agulation. Potato : Little or no growth. Acid but no gas within 48 hours from glucose, fructose, galactose, maltose, sucrose and mannitol. Acid after longer incubation from lactose, raffinose and glycerol. Indole is formed in small amounts. Nitrites not produced from nitrates. Aerobic. Is favored by increased CO2 tension. Will not grow anaerobi- cally. Optimum temperature 37 °C. Pathogenic for cattle and swine. A few cases reported in man. Rabbits and guinea pigs slightly susceptible to inocu- lation. Source and habitat : Usually isolated from the lesions of actinobacillosis of cattle. This condition is often clinically diagnosed as actinomycosis. Lesions found in soft tissues, usually lymph nodes, where granulomatous tumors are formed. Eventually these break down to form abscesses. 2. Actinobacillus actinomycetemcomi- tans Topley and Wilson. {Bacterium actinomycetem comitans Klinger, Cent, f. Bakt., I Abt., Orig., 1912, 198; Actino- bacillus actinomycetem comitans Topley and Wilson, Princip. of Bact. and Immun., 1, 1931, 256; Topley and Wilson, ibid., 2nd ed., 1936, 279.) From actinomycete, and Latin comitans, ac- companying. Description taken from Topley and Wilson {loc. cit.), Colebrook (Brit. Jour. Exp. Path., 1, 1920, 197), and Bayne-Jones (Jour. Bact., 10, 1925, 572). Cocco-bacilli : Rods 1.0 to 1.5 microns long, cocci 0.6 to 0.8 micron in diameter. Occurring in densely-packed masses. Non-motile. Gram-negative. Gelatin: No liquefaction. Agar colonies: Small, tough, adherent. Glucose agar: Growth thin, dry, gran- ular, hard, slightly yellow, adherent. Liquid gelatin or broth : At 37°C, nu- merous isolated, translucent granules, 0.5 to 1.0 mm in diameter, form along sides of tube. In a few days they fuse into a grayish-white mass, forming ring around tube and pellicle over surface. Later granules become opaque, grayish -white . Glucose broth: Turbid. Yellowish flakes. Milk: No growth. Potato : No growth. Acid but no gas from glucose and lac- tose. Not pathogenic for laboratory animals. No growth at 20°C. 558 MANUAL OF DETERMIXATIVE BACTERIOLOGY Aerobic, facultative. Distinctive character : Manner of growth in liquid gelatin. Source : Found in lesions of actinomy- cosis. Habitat : Presumably in actinomycotic lesions. 3. Actinobacillus actinoides (Smith) Topley and Wilson. ( Bacillus actinoides Th. Smith, Jour. Exp. Med., 28, 1918, 333; Actinomyces actinoides Bergey et al.. Manual, 1st ed., 1923, 346; Topley and Wilson, Princip. of Bact. and Immun., 1st ed., 1, 1931, 256.) From Greek, ray- like. Slender rods in tissues. In cultures may be bacillary or coccoid in form. Grows only under increased CO-, tension (so-called microaerophilic). Does not grow on ordinary agar or broth, except occasionally when transferred from more favorable media. Most characteristic growth on coagulated blood serum. Gelatin: No growth. Agar colonies : Very minute, pale, straw color. Agar slant : Best growth seen in water of condensatioia. Serial transfers on this medium generally fail. Broth: No growth. Litmus milk: No growth. Potato: No growth. Coagulated blood serum (cow) : Growth appears first in the condensation water. Appear as granules, consisting of cap- sular material in which bacillary forms are embedded. Surface mulberry-like because of club-like extensions of capsu- lar material. In stained preparations, the capsular material appears amorphous. Optimum temperature 37 °C. Microaerophilic. Not pathogenic for laboratory animals, except possibly the white rat in which a spontaneous chronic pneumonia occurs caused by an organism indistinguishable from this one. Experiments with rats by artificial inoculation have not been reported. Source : From lungs of calves suffering from chronic pneumonia. Habitat : Has not been recognized in nature except in pathological processes. FAMILY parvobactp:riaceae 559 * APPENDIX TO TRIBE PASTEURELLEAE. While the authors who describe the following new genus with its single species do not indicate its general relationships, it would appear to be as closely related to the species placed in Parrobacteriaccae as to those in anj' other family. It is therefore placed in this appendix pending a clarification of the situation. Genus A. Donovania Anderson, De Monbreun and Goodpasture. (Jour. Exp. Med., 81, 1945, 25.) Named for C. Donovan who first described the type species. Pleomorphic non-motile rods, exhibiting single or bipolar condensations of chromatin. Occur singly and in clusters. May be capsulated or non-capsulated. Gram -negative. Growth outside human body occurs only in the yolk, yolk sac or amniotic fluid of developing chick embryo or in a medium containing embryonic yolk. Pathogenic for man causing granulomatous lesions, particularly in the inguinal region. The type species is Donovania granulomatis Anderson, De Monbreun and Good- pasture . 1. Donovania granulomatis Anderson et al. (Epithelial cell parasites, Dono- van, Indian Med. Gaz., Jfi, 1905, 414; Donovan bodies, Dienst, Greenblatt and Sanderson, Jour. Inf. Dis., 62, 1938, 112; Donovan organism, Anderson, Science, 97, 1943, 560; Anderson, De Monbreun and Goodpasture, Jour. Exp. Med., 81, 1945, 25.) From M. L. granuloma, of granuloma. Pleomorphic rods 1 to 2 microns in length, with i-ounded ends, occurring singly and in clusters. Intracellular forms usually capsulated. Non-motile. Gram-negative. No growth on ordinary culture media. Chick embryo : Grows readilj^ in yolk, yolk sac and feebly in amniotic fluid of developing chick embryo. Embryonic yolk medium : Growth occurs. Distinctive characters : Capsulated forms readily demonstrated by means of Wright's stain as blue bacillary bodies surrounded by well-defined dense pink- ish capsules. Non-capsulated forms variable in morphology. Characteristic safety-pin forms may be demonstrated. Not pathogenic for the common experi- mental animals. Source : Granulomatous lesions of man. Habitat : Human lesions. The cause of granuloma inguinale. * Prepared by Dr. Orren D. Chapman, Syracuse Medical College, Syracuse, New York, March, 1946. 560 MANUAL OF DETERMINATIVE BACTERIOLOGY TRIBE II. BRUCELLEAE BERGEY, BREED AND MURRAY. (Preprint, Manual, 5th ed., October, 1938, vi.) Small, motile or non-motile rods or coccoids which grow on special media. There is a single genus Brucella. Genus I. Brucella Meyer and Shaw* (Jour. Inf. Dis., 27 , 1920, 173.) Named for Sir David Bruce, who first recognized the organism causing undulant fever. Short rods with many coccoid cells, 0.5 by 0.5 to 2.0 microns ; non-motile ; capsulated ; Gram-negative; gelatin not liquefied; neither acid nor gas from carbohydrates; urea utilized; parasitic, invading all animal tissues, producing infection of the genital organs, the mammary gland, the respiratory and intestinal tracts; pathogenic for various species of domestic animals and man. The type species is Brucella melitensis (Hughes) Meyer and Shaw. Key to the species of genus Brucella. I. Non-motile. A. Grow in special media containing basic fuchsin. 1. Grows in media containing thionin. 1. Brucella melitensis. 2. Does not grow in media containing thionin. 2. Brucella abortus. B. Does not grow in media containing basic fuchsin. 1. Grows in media containing thionin. 3. Brucella sui's. II. Motile. 4. Brucella bronchiseptica. Differential characters of the three closely related species of genus Brucella. Species Infec- tivity for guinea pigs Re- quires CO: for iso- lation H2S forma- tion *Glucose utilized Amino- nitio- gen utilized + + + + Growth in the presence of Thio- nin Basic fucbsin Brucella melitensis Brucella abortus + + + + + + 10 per cent 0 ++ 0 days ±1 2 4 + + + + + + + + + + 0 +++ + + + + + + Brucella suis 0 * All utilize glucose in shake cultures. 1. Brucella melitensis (Hughes) Meyer and Shaw. (Bruce, Practitioner, S9, 1887, 161; ibid., 40, 1888, 241; Kept. Army Med. Dept., London, 82, 1890, Append. No. 4, 465; streptococcus Mile- tensis (sic) Hughes, The Mediterranean Naturalist, 2, February 1, 1892, 325; Micrococcus melitensis Bruce, Ann. Inst. * Revised by Prof. I. F. Huddleson, Michigan State College, East Lansing, Michigan, December, 1942. FAMILY PARVOBACTERIACEAE 561 Past., 7, April, 1893, 289; Hughes, La Riforma Med., S, Aug. or Sept., 1893, 789 and Ann. Inst. Past., 7, Aug., 1893, 630; BacteriiDu melitense Ssiisawsi, Ztschr. f. Hyg., 70, 1912, 181; Meyer and Shaw, Jour. Inf. Dis., 27, 1920, 173; Bacillus melitensis Holland, Jour. Bact., 5, 1920, 219; Alcaligenes melitensis Bergey et al.. Manual, 1st ed., 1923, 235; Brucella meli- tensis var. melitensis Evans, U. S. Public Health Reports, 38, 1923, 1947.) From Latin, of Malta. Short ellipsoidal rods : 0.3 to 0.4 micron in length, occurring singly and in pairs, rarely in short chains. Non-motile. Non-acid-fast. Gram-negative. Gelatin colonies: Small, clear, entire. Gelatin stab: Slow growth. No lique- faction. Agar colonies: Small, circular, convex, amorphous, smooth, glistening, entire, bluish-green, grayish if R type. Agar slant : Growth slow, moist, honey- like, entire. After a week, the agar is turned brownish and crystals may ap- pear. Broth: After 10 days, moderate turbid- ity and grayish sediment. Reaction al- kaline, pH 8.0 or higher. Litmus milk: LTnchanged at 24 hours. Later becomes alkaline. Potato: Scant growth, grayish becom- ing brownish. Indole not formed. Nitrates reduced, often with complete disappearance of nitrite (Zobell and Meyer, Jour. Inf. Dis., 51. 1932, 99). Because of the latter fact, reports in the literature are apparently contradictor}'. Ammonia produced from urea. Growth enhanced on beef liver or tryptose agar of pH 6.8. Neither acid nor gas from carbohydrate media. Optimum reaction pH 7.4. Optimum temperature 37°C. No growth at 6° or at 45°C. Killed at 59°C. Aerobic. Distinctive characters : Requires no increased COo tension. Source : Isolated by Bruce (1887, loc. cit.) from the spleen in fatal cases of Malta fever. Habitat: Chief host the milch goat. The cause of undulant fever (brucellosis) in man and abortion in goats. IMay in- fect cows and hogs and be excreted in their milk. Infectious for all domestic animals. 2. Brucella abortus (Schmidt and Weis) Meyer and Shaw. (Bacillus of abortion. Bang, Ztschr. f. Thiermed., 1, 1897, 241 ; Bacterium abortus Schmidt and Weis, Bakterierne, 1901, 266; Bac- terium abortivum Chester, Man. Determ. Bact., 1901, 121 ; Corynebacterium abortus endemici Preisz, Cent. f. Bakt., I Abt., Orig., 33, 1902, 194; Bacillus abortus Evans, Jour. Wash. Acad. Sci., 5, 1915, 122; Meyer and Shaw, Jour. Inf. Dis., 27, 1920, 173; Alcaligenes abortus Bergey etal., Manual, 1st ed., 1923, 234; Brucella melitensis var. abortus Evans, Public Health Reports, 38, 1923, 1947.) From Latin abortus, an untimely birth. The morphological and cultural charac- ters are similar to those of Brucella melitensis with the following exceptions : Requires 10 per cent CO2 for isolation, becomes aerobic after several transfers; the browning of the medium in agar slant culture is less marked; S cultures can be difl'erentiated from Brucella melitensis, but not from Brucella suis, by the ag- glutinin absorption test. Source : From the genital organs and milk of infected cattle and from blood in human cases of undulant fever. Habitat: Chief host the milch cow. The cause of infectious abortion in cattle. The same effects are produced in mares, sheep, rabbits and guinea pigs, and all domestic animals except hogs. Causes undulant fever (brucellosis) in man. 3. Brucella suis Huddleson. (Organ- ism resembling Bacillus abortus. Anony- mous, U. S. D. A. Ann. Rept. Secy. Dept., Rept. of Chief Bur. Animal Ind., 562 MANUAL OF DETERMINATIVE BACTERIOLOGY 1914, 86 (30); authorship established by Traum in North Amer. Vet., 1, No. 2, 1920; described as Bacillus abortus by Good and Smith, Jour. Bact., 1, 1916, 415; Huddleson, Undulant Fever Sym- posium, Amer. Pub. Health Assoc, (Oct., 1928) 1929, 24; also Mich. Agr. Exp. Sta. Tech. Bull. 100, 1929, 12; Brucella melitensis var. suis Hardy, Jordan, Borts and Hardy, Public Health Reports, 4S, 1930, 2433; Bacillus abortus suis Meyer, Amer. Jour. Pub. Health, 21, 1931, 503.) From Latin, of swine. The morphological and cultural charac- ters are similar to those of Brucella melitensis. S cultures of Brucella suis can be dif- ferentiated from S cultures of Brucella melitensis, but not from S cultures of Brucella abortus, by the agglutinin absorption test. Source : From urinogenital and many other organs of swine. Habitat : Chief host the hog. Causes abortion in swine and undulant fever (brucellosis) in man. Also infectious for horses, dogs, cows, monkeys and laboratory animals. The ditferentiation of the above species of Brucella by the bacteriostatic action of dyes depends upon the medium used. When tryptose agar (Difco) is used, basic fuchsin and thionin should be used in a final dilution of 1:100,000. There are several forms of the R and mucoid phases of Brucella spp. (Huddle- son, Amer. Jour. Vet. Res., 7, 1946, 5). The true R type differs from the S type in its lack of pathogenicity, its antigenic properties, its susceptibility to aggluti- nation by exposure of suspensions to heat and to basic dyes in concentration of 1 : 2000, and colonial appearance. The mucoid phases differ antigenically, mor- phologically and culturally. Colonies on agar are spherical or flat, regular in contour, grayish to mucoid in appear- ance. Suspensions are not agglutinated by heat or dyes, or always by special agglutinating serums. There is no change in their growth characteristics on media containing either basic fuchsin or thionin. 4. Brucella bronchiseptica (Ferry) Topley and Wilson. (Ferry, Amer. Vet. Rev., 37, 1910, 499; also see McGowan, Jour. Path., 15, 1911, 372; Bacillus bron- chicanis Ferry, Jour. Inf. Dis., 8, 1911, 402; Bacillus bronchisepticus Ferry, Amer. Vet. Rev., 41, 1912, 79; Bacterium bronchisepticus Evans, Jour. Inf. Dis., 18, 1916, 578; Bacterium bronchicanis Holland, Jour. Bact., 5, 1920, 221 ; Alcali- genes bronchisepticus Bergey et al., Manual, 1st ed., 1923, 234; Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 508.) Latinized, disease of the bronchial tubes. Evans {loc. cit., 593) regards Bacterium bronchisepticus as related to Bacterium abortus morphologically, culturally, bio- chemically and serologically. Short slender rods: 0.4 to 0.5 by 2.0 microns, usually occurring singly, some- times in pairs and chains. Motile with 4 to 6 peritrichous flagella (Topley and Wilson). Gram-negative. Gelatin colonies : Similar to those on agar. Gelatin stab : Slow filiform growth. No liquefaction. Agar colonies: Small, opaque, white, slightly raised, porcellaneous, entire. Agar slant : Growth moderate but more luxuriant than in Brucella melitensis, filiform, slightly raised, smooth, opales- cent, lustrous, moist, entire. Broth: Turbid, with thin, gray pellicle and ropy sediment. Musty odor de- velops. Litmus milk: Alkaline. No coagula- tion. Potato : Growth fairly abundant, brownish, glistening, moist, sticky. Me- dium is darkened. Indole not formed. Nitrites often produced from nitrates (Topley and Wilson). FAMILY PARVOBACTERIACEAE 563 No acid or gas from glucose, sucrose, lactose, maltose or mannitol. No H2S produced (Toplej' and Wilson). Catalase positive (Topley and Wilson). Ammonia formed from urea and as- paragine. Optimum temperature 37°C. Ivilled in twenty minutes at 55°C. Aerobic, facultative. Source : From dogs affected with dis- temper. Habitat: Causes acute, often fatal, pneumonia in dogs generally as a second- ary invader in distemper. Also patho- genic for cats, rabbits, guinea pigs, fer- rets, white rats and monkeys. Some- times occurs in man. Appendix: The following are recorded in the literature discussing this genus: Brucella evansi Pacheco (Revista da Sociedade Paulista de Med. Vet., S, 1933, 9) is a name applied to a group of thirteen cultures referred to by Evans (Jour. Inf. Dis., 23, 1918, 354) as abortus- like bacteria although she definitelj' in- dicates that these cultures do not agree with each other in their biochemical characteristics (loc. cit., Table 4, p. 361). The binomials Brucella -paramelitensis , Brucella paraabortus and Brucella para- suis have been used for inagglutinable strains of these three species which are, according to Topley and Wilson (Princip. Bact. and Immun., 2nd ed., 1936, 632), now known to be merely rough variants, not deserving to be so named. Micrococcus paramelitensis Negr^ and Raymond. (Compt. rend. Soc. Biol., Paris, 72, 1912, 791 and 1052.) Micrococcus pseudomelitensis Sergent and Zammitt, 1908. Exact reference not 564 MANUAL OF DETERMIXATIVE BACTERIOLOGY TKIBE III. BACTEROIDEAE TRIE. NOV. Motile or non-motile rods without endospores. May or may not require enriched culture media. Obligate anaerobes. Gram -negative. Key to the genera of tribe Bacteroideae. I. Cells with rounded ends. Genus I. Bacteroides, p. 564. II. Cells with pointed ends. Genus II. Fusohacterium, p. 581. Genus I . Bacteroides Castellani and Chalmers.* (Man. Trop. Med., 3rd ed., 1919, 959.) Characters as for the tribe. From Greek, like a rod. The type species is Bacteroides fragilis (Veillon and Zuber) Castellani and Chalmers. Note : The descriptions have been taken largely from Weinberg et al. (Les Microbes Anaerobies, Paris, 1937, 658); Prevot (Ann. Inst. Past., 60, 1938, 285); Hauduroy, Ehringer, Urbain, Guillot and Magrou (Diet. Bact. Path., Paris, 1937, 51); and Eg- gerth and Gagnon (Jour. Bact., 25, 1933, 389). Because cultures of many of these organisms have not been subjected to critical study with identical tests and media, it is difficult to know how many should be considered as distinct species, and the present arrangement must be considered as tentative. The key, of necessity, has been drawn up from recorded characters which appeared useful for. the purpose and these on further study may prove to be inadequate. Key to the species of genus Bacteroides. I. Not requiring enriched media. A. Gas formed from proteins. 1. Hydrogen sulfide not produced, a. Non-motile. aa. Motile. 2. Hydrogen sulfide produced, a. Indole not formed. b. Very pleomorphic. 1. Bacteroides fragilis. 2. Bacteroides serpens. 3. Bacteroides funduliformis. bb. Not markedly pleomorphic. 4. Bacteroides siccus. aa. Indole formed. b. Gelatin liquefied. 5. Bacteroides coagulans. * Completely revised by Dr. T. E.Roy, Bacteriologist to the Hospital for Sick Chil- dren, Toronto, Ontario, Canada and Dr. C. D. Kelly, Assistant Professor of Bacteriol- ogy, McGill University, Montreal, P. Q., Canada, December, 1938; rearranged, Decem- ber, 1945. FAMILY PARVOBACTERIACEAE 565 bb. Gelatin not liquefied. c. No acid from lactose and maltose. 6. Bacteroides varius. cc. Acid from lactose and maltose. d. Acid from sucrose. No acid from glycerol. 7. Bacteroides inaequalis. dd. No acid from sucrose. Acid from glycerol. 8. Bacteroides insolitus. B. No gas formed from proteins. 1. Indole not formed. a. Hydrogen sulfide not formed, b. No acid from lactose. 9. Bacteroides vescus. bb. Acid from lactose. c. No acid from salicin. 10. Bacteroides exiguus. cc. Acid from salicin. 11. Bacteroides uncatus. aa. Hydrogen sulfide formed. b. No acid from salicin. Acid from arabinose. c. Gelatin liquefied. 12. Bacteroides vulgatus. cc. Gelatin not liquefied. 13. Bacteroides incommunis. bb. Acid from salicin. No acid from arabinose. 14. Bacteroides distasonis. bbb. No acid from salicin or arabinose. c. Acid from sorbitol. 15. Bacteroides tumidus. cc. No acid from sorbitol. 16. Bacteroides convexus. 2. Indole formed. a. No acid from salicin or arabinose. 17. Bacteroides ovatus. aa. Acid from salicin and arabinose. b. No acid from mannitol. c. No acid from rhamnose. 18. Bacteroides uniformis. cc. Acid from rhamnose. d. Not capsulated. 19. Bacteroides thetaiotaomicron. dd. Capsulated. bb. Acid from mannitol. II. Requiring an enriched medium. A. Producing a black pigment. B. Not producing pigment. 20. Bacteroides variabilis. 21. Bacteroides gulosus. 22. Bacteroides melaninogenicus. 23. Bacteroides caviae. 566 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Bacteroides fragilis (Veillon and Zuber) Castellani and Chalmers. (Bacillus fragilis Veillon and Zuber, Arch. Med. Exp. et Anat. Path., 10, 1898, 870; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 959; Fusijormis fragilis Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 302; Ristella fragilis Prevot, Ann. Inst. Past., 60, 1938, 290.) From Latin fragilis, fragile. Rods with rounded ends, staining more deeply at the poles, occurring singly and in pairs. Non-motile. Gram- negative. Gelatin : No liquefaction ; small amount of gas. Agar colonies: Small, graj^ irregular. Broth: Turbid. Indole not formed. Hydrogen sulfide not formed. Litmus milk: No coagulation. Slight amount of gas. Nitrites not produced from nitrates. Acid from fructose, maltose, sucrose, galactose, glucose and arabinose. Some strains produce acid from lactose (Wein- berg et al., Les Microbes Ana^robies, 1937, 720). Anaerobic. Optimum temperature 37 °C. Pathogenicity: Some strains produce subcutaneous abscesses in rabbits, guinea pigs or mice. Source and habitat : From acute appen- dicitis, pulmonary gangrene, abscesses of the urinary tract, and septicaemias in man. 2. Bacteroides serpens (Veillon and Zuber) Hauduroy et al. {Bacillus serpejis Veillon and Zuber, Arch. Med. Exp. et Anat. Path., 10, 1898, 870; Bacillus radiiformis Rist and Guillemot, Arch. Mdd. Exp. et Anat. Path., 1904; Haudu- roy et al.. Diet. d. Bact. Path., 1937, 74; Zuherella serpens Prevot, Ann. Inst. Past., 60, 1938, 293.) From Latin ser- pens, creeping. Rods : Thick, with rounded ends, oc- curring singly, in pairs, or in short chains. Motile. Gram-negative. Gelatin: Slow liquefaction, with gas. Agar colonies: Punctiform. Deep agar colonies: Small colonies in 48 hours, ray-like growth later. Gas produced. Broth : Turbid, then flocculent growth ; some gas with foul odor. Hj'drogen sulfide not formed. Litmus milk: Acidified and coagulated in six daj'S, with no digestion. Acid from fructose, galactose, maltose and lactose. Coagulated egg white and serum not liquefied. Anaerobic. Optimum temperature 37°C. Experimental pathogenicity : Some strains produce abscesses in rabbits, guinea pigs and mice. Source and habitat : Acute appendi- citis, mastoiditis, pulmonary gangrene, l)ile tract of dog, and sea water. 3. Bacteroides funduliformis (Halle) Bergey et al. {Bacilhis funduliformis Hall^, Inaug. Diss., Paris, 1898; Bacillus thetoides Rist, The«e de Paris, 1898; Bergey et al., Manual, 3rd ed., 1930, 373; Spherophorus funduliformis Prevot, Ann. Inst. Past., 60, 1938, 298.) From Latin funduliformis, sausage -shaped . Rods: 1.5 to 3.0 microns long in pus, often spindle-shaped. Extremely pleo- morphic in culture media, showing irreg- ular filamentous and branching forms. Non-motile. Gram-negative. Gelatin: Not liquefied. Deep agar colonies: Lenticular, with some gas and foul odor. Broth: Flocculent growth. Glucose broth : Rapid growth with gas and foul odor. Indole not formed; although sometimes found in old cultures. Hydrogen sulfide is formed in small amounts . Litmus milk : Acid and coagulation by some strains. FAMILY PARVOBACTERIACEAE 567 Acid and gas from fructose, glucose and maltose. Some strains ferment man- nitol, sucrose and lactose. Anaerobic. Optimum temperature 37 °C. Experimental pathogenicity : Some strains are pathogenic for rabbits and guinea pigs Init not for white rats and mice. Source and habitat: Female genitalia, urinary infections, puerperal infections, acute appendicitis, otitis, pulmonary gangrene, liver abscesses, septicaemias and intestinal tract. 4. Bacteroides siccus Eggerthand Gag- non. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 410; Spherophorus siccus Prevot, Ann. Inst. Past., 60, 1938, 299.) From Latin siccus, dry. Short, thick rods: About 1.0 micron long. In glucose broth they are coccoid and often grow in short chains. Non- motile. Gram-negative. Gelatin: Xot liquefied. Blood agar colonies: Elevated, dry, difficult to emulsify, 1.0 to 1.5 mm in diameter. Broth: Growth occurs as a powdery sediment with a clear supernatant fluid. Indole not formed. Hj'drogen sulfide is formed. Milk: Unchanged. Nitrites not produced from nitrates. Acid but no gas from fructose. No acid or gas from glucose, glycerol, mannitol, sorbitol, arabinose, saliein, trehalose, amygdalin, cellobiose, glj'^cogen, rham- nose, xylose or lactose. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters: Gas is formed in small amounts from peptone. Phenol red and brom cresol purple are de- colorized in meat infusion broth. Source : Two strains isolated from human feces. Habitat: Probably intestinal canal of mammals. 5. Bacteroides coagulans Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 409 ; Pasteur ella coagulans Prevot, Ann. Inst. Past., 60, 1938, 292.) From Latin coagulans, coagulating. Rods : 0.5 to 2.0 microns long. Bipolar staining. Non-motile. Gram-negative. Gelatin: Liquefied in 8 to 12 days. Blood agar colonies: Soft, transparent, 0.5 mm in diameter. Broth : Diffuse growth. Indole is formed. Hydrogen sulfide is formed. Milk: Coagulated in 8 days without acid production. The coagulum partly redissolves after 3 to 4 weeks. Nitrites not produced from nitrates. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters : No acid or gas from carbohydrates. A small amount of gas is formed from peptone. Phenol red and brom cresol purple are de- colorized in a meat infusion broth. Source : One strain isolated from human feces. Habitat : Probably intestinal canal of mammals. 6. Bacteroides varius Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 409; Spherophorus varius Prevot, Ann. Inst. Past., 60, 1938, 299.) From Latin varius, diverse. Rods : 1 .0 to 3.0 microns long. Staining uneven. Non-motile. Gram-negative. Gelatin: Not liquefied in 45 daj's. Blood agar colonies: Very flat cones, 2.0 to 3.0 mm in diameter. Broth: Diffusely clouded. Indole is formed. Hydrogen sulfide produced. Milk: Not acidified or coagulated. Nitrites not produced from nitrates. Acid and gas from fructose, galactose, glucose and mannose. No acid or gas from esculin, amygdalin, arabinose, cello- biose, dextrin, glycerol, glycogen, inulin, lactose, maltose, mannitol, melezitose, 568 MANUAL OF DETERMINATIVE BACTERIOLOGY raffinose, rhamnose, salicin, sorbitol, starch, sucrose, trehalose or xylose. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters : Gas is formed from peptone. Brom cresol purple and phenol red are decolorized in a meat in- fusion broth. Source : Two strains isolated from hu- man feces. Habitat : Probably intestinal canal of mammals. 7. Bacteroides inaequalis Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 407; Spherophorus inae- qualis Pr^vot, Ann. Inst. Past., 60, 1938, 298.) Fi"om Latin inaequalis, unequal. Rods: Wide variation in size and form. Marked pleomorphism on blood agar. Non-motile. Gram-negative. Gelatin: Not liquefied in 45 days. Blood agar colonies: Pin-point in size. Broth: Diffusely clouded. Indole is formed. Hydrogen sulfide is produced. Milk: Acidified but not coagulated. Nitrites not produced from nitrates. Acid but no gas from esculin, amyg- dalin, arabinose, fructose, galactose, glucose, lactose, maltose, mannose, raffi- nose, salicin, sucrose and xylose. No acid or gas from cellobiose, dextrin, glycerol, glycogen, inulin, mannitol, melezitose, rhamnose, sorbitol, starch and trehalose. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters : Forms small amount (5 per cent in Smith tube) of gas from peptone water in the complete absence of carbohydrates. None of this gas is absorbed by alkali. Rapidly de- colorizes brom cresol purple and phenol red in meat infusion broth; slowly or not at all in peptone water. Source : One strain isolated from human feces. Habitat: Probably intestinal canal of mammals. 8. Bacteroides insolitus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 408; Ristella insolita Prevot, Ann. Inst. Past., 60, 1938, 291.) From Latin insolitus, uncommon. Short thick rods: 1.0 to 2.0 microns long. Often slender, curved, 2.0 to 3.0 microns long. Non-motile. Gram- negative. Gelatin: Not liquefied in 45 days. Blood agar colonies: Minute, trans- parent. Broth: Heavy, diffuse growth. Indole is formed. Hydrogen sulfide is formed. Milk: Acidified and coagulated in 30 to 35 days. Nitrites not produced from nitrates. Acid but no gas from fructose, galac- tose, glucose, glycerol, lactose, maltose and mannose. No acid or gas from es- culin, amygdalin, arabinose, cellobiose, dextrin, glycogen, inulin, mannitol, melez- itose, raffinose, rhamnose, salicin, sor- bitol, starch, sucrose, trehalose and xylose . Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters: Brom cresol purple and phenol red are rapidly de- colorized in a meat infusion broth. A small amount of gas is formed from pep- tone . Source : One strain isolated from human feces. Habitat : Probably intestinal canal of mammals. 9. Bacteroides vescus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 406; Fusiformis vescus Prdvot, Ann. Inst. Past., 60, 1938, 300.) From Latin vescus, small or weak. Slender, pointed rods: 1.0 to 2.0 microns long, sometimes slightly curved. FAMILY PARVOBACTERIACEAE 569 Bipolar staining. Non-motile. Gram- negative. Gelatin: Liquefied in 8 to 25 days. Blood agar colonies : Very minute and transparent. Broth : Diffusely clouded. Indole not formed. Hydrogen sulfide not produced. Milk: Neither acidified nor coagulated. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas from cellobiose (in 30 days), dextrin, glucose, maltose, man- nose and rhamnose. No acid or gas from esculin, amygdalin, arabinose, galactose, mannitol, melezitose, raffinose, salicin, sorbitol, starch, sucrose, trehalose, xylose, glycerol, glycogen, inulin, lactose or fructose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : One strain isolated from hu- man feces. Habitat : Probably intestinal canal of mammals. 10. Bacteroides exiguus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 407; Ristclla exigua Prd- vot, Ann. Inst. Past., 60, 1938, 292.) From Latin exiguus, small and narrow. Very small slender rods: 0.5 to 1.0 micron long, occurring singly and in pairs. Non-motile. Gram-negative. Gelatin: Liquefied in 16 to 20 days. Blood agar colonies : These are of two types. One is pin-point in size, the other is large, gray, moist, 1.0 to 1.5 mm in diameter. Broth: Diffusely clouded. Indole not formed. Hydrogen sulfide not formed. Milk : Acidified and may or may not be coagulated in 35 to 40 days. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas from fructose, galac- tose, glucose, lactose, maltose, mannose, sucrose and trehalose. One strain fer- ments raffinose. No acid or gas from esculin, amygdalin, arabinose, cellobiose, dextrin, glycerol, glycogen, inulin, man- nitol, melezitose, rhamnose, salicin, sor- bitol, starch or xylose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : Two strains isolated from human feces. Habitat : Probably intestinal canal of mammals. 11. Bacteroides uncatus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 404; Ristella uncata Pr^vot, Ann. Inst. Past., 60, 1938, 291.) From Latin uncatus, hooked at the tip. Rods : Extreme variations in size and form. The ordinary length is 5.0 to 8.0 microns. Curved and hooked forms common. Non-motile. Gram-negative. Gelatin: Liquefied in 16 days. Blood agar colonies : Very minute and transparent. Broth: Turbid, growth is slow and light. Indole not formed. Hydrogen sulfide not formed. Milk : Not acidified or coagulated. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas after 8 to 30 days of incubation from dextrin, fructose, galac- tose, glucose, lactose, maltose, raffinose, rhamnose, salicin, starch and sucrose. No acid from esculin, amygdalin, arabi- nose, cellobiose, glycerol, glycogen, in- ulin, mannitol, mannose, melezitose, sorbitol, trehalose or xylose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : One strain isolated from human feces. Habitat : Probably intestinal canal of mammals. 12. Bacteroides vulgatus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. 570 MANUAL OF DETERMINATIVE BACTERIOLOGY Bact., 25, 1933, 401; Pasteurella vulgata Pr^vot, Ann. Inst. Past., 60, 1938, 292.) From Latin vulgatus, common. Oval rods : 0.7 to 2.5 microns long, usu- ally occurring singly, sometimes in pairs. One strain formed filaments 10 microns long. Stain solidlj'', some strains show bipolar staining. Morphol- ogy very variable in glucose broth. Non-motile. Gram-negative. Gelatin: Liquefied in 4 to 20 days by all but one strain. Blood agar colonies: Soft, translucent, grayish, elevated, 1.5 to 2.0 mm in diam- eter. Half of the strains are hemolytic. Broth: Heavy and diffuse growth. Indole not formed. Hydrogen sulfide is formed. Milk : Acidified. Coagulated by some strains in 5 to 25 days. Nitrites not produced from nitrates. Acid and a small amount of gas from arabinose, dextrin, fructose, galactose, glucose, glycogen, inulin, lactose, maltose, mannose, raffinose, rhamnose, starch, sucrose and xylose. Seven strains fermented esculin. No acid or gas from amygdalin, cellobiose, glycerol, mannitol, melezitose, salicin, sorbitol, trehalose, dulcitol, erythritol or inositol. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters: Does not form indole; does not produce gas from pep- tone. This is the commonest species found in the feces of adults. Differs from Bacteroides incormnunis in that it does not ferment amygdalin and cello- biose, but does ferment glycogen and starch. Liquefies gelatin. Source : Thirty-eight strains isolated from human feces. Habitat : Probably intestinal canal of mammals. 13. Bacteroides incommunis Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 402; Ristella incom- munis Prdvot, Ann. Inst. Past., 60, 1938, 291.) From Latin incommunis, not common. Rods: 0.5 to 1.5 by 1.0 to 3.0 microns, occurring singly. Stain solidly. Non- motile. Gram-negative. Gelatin: Not liquefied. Blood agar colonies : Elevated, slightly yellowish, 1 mm in diameter. One strain formed soft colonies; the other was stringy when emulsified. Broth: Growth is diffuse. Indole not formed. Hydrogen sulfide is formed. Milk: Acidified but not coagulated; coagulates promptly on boiling. Nitrites not produced from nitrates. Peptone : No gas . Acid and a small amount of gas from amygdalin, arabinose, cellobiose, dextrin, fructose, galactose, glucose, inulin, lac- tose, maltose, mannose, raffinose, rham- nose, sucrose and xylose. One strain fermented glycogen and starch. No action on esculin, glycerol, mannitol, melezitose, salicin, sorbitol or trehalose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : Two strains isolated from hu- man feces. Habitat : Probably intestinal canal of mammals. 14. Bacteroides distasonis Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 403; Ristella distasonis Pr^vot, Ann. Inst. Past., 60, 1938, 291.) Named for Distaso, Roumanian bac- teriologist. Rods: 0.5 to 0.8 by 1.5 to 2.5 microns, occurring singly. Staining solidly and having rounded ends. Some strains show a few bacilli 5.0 to 8.0 microns long. Non-motile. Gram-negative. Gelatin: Not liquefied by 16 strains. The remaining 4 liquefied gelatin in 35 to 50 days. Blood agar colonies : Soft, grayish, elevated colonies, 1.0 to 1.5 mm in diam- eter. Two strains markedly hemolytic. FAMILY PARVOBACTERIACEAE 571 Broth: Growth is diffuse. Indole not formed. Hydrogen sulfide is produced. Milk: Acidified. All but 4 strains coagulate milk. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas from amygdalin, cellobiose, dextrin, fructose, galactose, glucose, inulin, lactose, maltose, man- nose, melezitose, raffinose, rhamnose, salicin, sucrose, trehalose and xylose. Fifteen strains ferment esculin. Fifteen strains slowly ferment starch. No acid or gas from arabinose, glycogen, glycerol, mannitol or sorbitol. Non-pathogenic for white mice and rab- bits. Anaerobic. Distinctive characters: Usually fails to liquefy gelatin. Fails to ferment arabinose . Source : Twenty strains isolated from human feces. Habitat : Probably intestinal canal of mammals. 15. Bacteroides tumidus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 2o, 1933, 405; Ristella tumida Prevot, Ann. Inst. Past., 60, 1938, 292.) From Latin tumidus, swollen. Small, thick oval rods: 1.0 to 1.5 microns long and occurring singly. The staining is solid. On glucose broth many swollen forms with irregular staining from 1.0 to 4.0 by 1.5 to 10 microns. The bodies of these swollen forms are usually very pale, with only the ends staining. Non-motile. Gram-negative. Gelatin: Liquefied in 12 to 20 daj^s. Blood agar colonies: Soft, grayish, elevated colonies, 1 mm in diameter. Broth: Heavy, diffuse growth. Indole not formed. Hydrogen sulfide is produced. Milk: Acidified but not coagulated. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas from dextrin, fruc- tose, galactose, glucose, glycogen, inulin, lactose, maltose, mannose, raffinose, sor- bitol, starch and sucrose. No acid or gas from esculin, amygdalin, arabinose, cellobiose, glycerol, mannitol, melezitose, rhamnose, salicin, trehalose or xylose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : Four strains isolated from human feces. Habitat : Probably intestinal canal of manmials, 16. Bacteroides convexus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 406; Pasteurella convexa Prevot, Ann. Inst. Past., 60, 1938, 292.) From Latin convexus, convex. Thick, oval rods: 0.8 to 1.5 microns long, occurring singly or in pairs. In glucose broth, the rods are usually 2.0 to 3.0 microns long. Non-motile. Gram- negative. Gelatin : Liquefied in 20 to 30 days. Blood agar colonies: Elevated, grayish, somewhat opaque colonies, 1.0 to 1.5 mm in diameter. Broth: Heavy diffuse growth. Indole not formed. Hydrogen sulfide is produced. Milk: Acidified and coagulated in 4 days. Nitrites not produced from nitrates. Peptone : No gas. Acid and a small amount of gas from esculin, amygdalin, cellobiose, dextrin, fructose, galactose, glucose, glycogen, inulin, lactose, maltose, mannose, raffi- nose, starch, sucrose and xylose. No acid or gas from arabinose, glycerol, mannitol, melezitose, rhamnose, salicin, sorbitol or trehalose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : Five strains isolated from human feces. Habitat: Probably intestinal canal of mammals. 572 MANUAL OF DETERMINATIVE BACTERIOLOGY 17. Bacteroides ovatus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 405; Pasteurella ovata Pr^vot, Ann. Inst. Past., 60, 1938, 292.) From Latin ovatus, egg-shaped. Small oval rods: 0.5 to 1.0 by 1.0 to 2.0 microns, occurring singly. Stains solidly. Non -motile. Gram-negative. Gelatin: Liquefied in 4 days. Blood agar colonies: Soft, grayish, elevated colonies, 1.0 to 1.5 mm in diam- eter. Broth: Diffuse, heavy growth. Indole is formed. Hydrogen sulfide is produced. Milk : Acidified and coagulated in 4 days. Nitrites not produced from nitrates. Peptone : No gas. Acid and a small amount of gas from esculin, amygdalin, cellobiose, dextrin, fructose, galactose, glucose, glj^cogen, inulin, lactose, maltose, mannose, rafR- nose, rhamnose, starch, sucrose and xylose. No acid or gas from arabinose, glycerol, mannitol, melezitose, salicin, sorbitol or trehalose. Non-pathogenic for white mice and rabbits. Anaerobic. Source : One strain isolated from human feces. Habitat : Probably intestinal canal of mammals. 18. Bacteroides uniformis Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 25, 1933, 400; Ristella uniformis Prdvot, Ann. Inst. Past., 60, 1938, 291.) From Latin uniformis, of a single form. Small rods: 0.8 to 1.5 microns long, occurring singly, with rounded ends. Stain heavier at poles and around periph- ery. Non-motile. Gram-negative. Gelatin : Liquefied by two strains in 15 to 40 days. Six strains did not liquef}^ Blood agar colonies: Transparent, soft, elevated, 0.5 to 0.75 mm in diameter. Broth: Diffuse growth. Indole formed. Hydrogen sulfide produced slowly or not at all. Milk : Acidified and coagulated in 8 to 12 days. Nitrites not produced from nitrates. Peptone : No gas. Acid but no gas from esculin, amyg- dalin, arabinose, cellobiose, dextrin, fruc- tose, galactose, glucose, glycogen, inulin, lactose, maltose, mannose, melezitose, raffinose, salicin, starch, sucrose, treha- lose and xylose. No acid or gas from glycerol, mannitol, rhamnose, sorbitol, dulcitol, erythritol or inositol. Non-pathogenic for white mice and rabbits. Anaerobic. Distinctive characters: Forms indole. Resembles Bacteroides vulgatus. Source: Eight strains isolated from human feces. Habitat : Probably intestinal canal of mammals. 19. Bacteroides thetaiotaomicron (Distaso) Castellani and Chalmers. {Bacillus thetaiotaomicron Distaso, Cent, f. Bakt., I Abt., Grig., 62, 1912, 444; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 960; Spherocillus thetaiotaomicron Pr^vot, Ann. Inst. Past., 60, 1938, 300.) The combination theta, iota and omicron is used because the pleomorphic rods have the shape of these Greek letters. Description taken from Distaso {loc. cit.). More complete description will be found in Eggerth and Gagnon (Jour. Bact., 25, 1933, 399). Short, plump to oval rods. Stain solidly or only at poles. Sometimes with bar causing organism to resemble Greek letter theta. Motile (Distaso). Non- motile (Eggerth and Gagnon). Gram- negative. Gelatin: No liquefaction. Glucose agar colonies: Large, trans- parent, entire. Sometimes form gas bubbles. Broth: Turbid. FAMILY PARVOBACTERIACEAE 573 Egg albumen broth : Albumen not attacked. Indole is formed. Hydrogen sulfide produced (Eggerth and Gagnon). Litmus milk: Acid, coagulated. Curd shrinks with expulsion of turbid whey. Nitrates not recorded (Distaso). Ni- trites not produced from nitrates (Eg- gerth and Gagnon). Peptone : No gas (Eggerth and Gag- non). Acid and gas from esculin, amygdalin, arabinose, fructose, inulin, lactose, cello- biose, dextrin, galactose, glucose, glj'co- gen, maltose, mannose, melezitose, raflinose, rhamnose, salicin, starch, su- crose, trehalose and xylose. Four strains fail to produce gas from any sugar. No acid or gas from glycerol, mannitol or sorbitol (Eggerth and Gagnon). Anaerobic . Distinctive characters : Resembles Bncteroides variabilis but is not cap- sulated, does not liquefy gelatin, usually forms gas from sugars, and ferments melezitose and trehalose. Differs from Bacteroides unifonnis in morphology, forming gas from sugars and in ferment- ing rhamnose (Eggerth and Gagnon). Source : Isolated frequently from human feces. Habitat : Intestinal canal of mammals (common ) . 20. Bacteroides variabilis (Distaso) Castellani and Chalmers. {Bacillus variabilis Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 441; Castellani and Chalmers, jVIan. Trop. Med., 3rd ed., 1919, 960; Capsularis variabilis Prevot, Ann. Inst. Past., ffO, 1938, 293.) From Latin variabilis, variable. Short rods, with rounded ends, occur- ring singly. Some long flexuous rods. Capsulated. Non-motile. Gram- negative . Gelatin: No growth on plain gelatin (Distaso); liquefaction (Eggerth and Gagnon, Jour. Bact., 25, 1933, 400). Blood agar colonies : Smooth, glistening, elevated and ver^' mucoid, about 1.0 mm in diameter. Broth: Diffuse growth. Indole is formed. Hj-drogen sulfide is formed. Litmus milk: Unchanged (Distaso); acidified and some strains coagulating in 25 to 35 days (Eggerth and Gagnon). Nitrites not produced from nitrates (Eggerth and Gagnon). Peptone : No gas. Acid and gas from glucose, lactose and sucrose (Distaso) . Acid and no gas from esculin. amygdalin, arabinose, cello- biose, dextrin, fructose, galactose, glycogen, inulin, lactose, glucose, maltose, mannose, raffinose, rhamnose, salacin. starch, sucrose and .xylose. No acid or gas from glycerol, mannitol, melezitose, sorbitol or trehalose (Eg- gerth and Gagnon). Non-pathogenic for white mice and rabbits. Anaerobic. Optimum temperature 37°C. Distinctive characters: Capsulated. Source : Isolated from human feces by Distaso, and by Eggerth and Gagnon (8 strains). Habitat : Probably intestinal canal of mammals. 21. Bacteroides gulosus Eggerth and Gagnon. (Eggerth and Gagnon, Jour. Bact., 2-5, 1933, 398; Spherophoriis gulosus Prevot, Ann. Inst. Past., 60, 1938, 298.) From Latin gulosus, gluttonous. Small oval rods: 0.8 to 1.0 by 1.0 to 2.0 microns, staining deeper around periph- ery. Non-motile. Gram-negative. Gelatin: Liquefied in 2 to 3 weeks. Blood agar colonies: Soft, gray, entire, elevated, 2 mm in diameter. Broth: Heavy and diffuse growth. Indole formed. Hj'drogen sulfide is formed. Milk : Acidified and coagulated in 4 to 20 days. Nitrites not produced from nitrates. 574 MANUAL OF DETERMINATIVE BACTERIOLOGY Acid and a very small amount of gas from esculin, amygdalin, arabinose, cel- lobiose, dextrin, fructose, galactose, glycogen, inulin, lactose, glucose, mal- tose, mannitol, mannose, melezitose, raffinose, rhamnose, salicin, sorbitol, starch, sucrose, trehalose and xylose. Sorbitol and mannitol require 2 to 3 weeks for fermentation. Neither acid nor gas from glycerol, dulcitol, erj'thritol or inositol. Peptone : No gas. Non-pathogenic for white mice and rabbits. Anaerobic. Source : Seven strains isolated from human feces. Habitat : Probably intestinal canal of mammals. 22. Bacteroides melaninogenicus (Oli- ver and Wherry) Roy and Kelly. {Bac- terium melaninogenicum Oliver and Wherry, Jour. Inf. Dis., 28, 1921, 341; Hemophilus melaninogenicus Bergey et al., Manual, 3rd ed., 1930, ZU; Ristella melaninogenica Pr^vot, Ann. Inst. Past., 60, 1939, 290; Roy and Kelly, in Manual, 5th ed., 1939, 569.) From Greek, black- producing. Description taken from Oliver and Wherry {loc. cit.) and Burdon (Jour. Inf. Dis., 42, 1928, 161). Rods: 0.8 by 1.0 to 3.0 microns. Non- motile. Gram-negative. Serum gelatin stab: Dense flocculent growth at 37°C. No liquefaction. Blood agar slant: Confluent, black, dry layer. The blood is disintegrated in one to two weeks forming melanin. The medium becomes colorless. Sodium phosphate broth: Turbid. Litmus milk : Slow acidification but no coagulation. Blood serum slant : Fairly luxuriant, white, moist layer. Acid from fructose, glucose, lactose, maltose, sucrose and mannitol. No acid from galactose. Non-pathogenic for rabbits, guinea pigs and white mice (Burdon). Anaero- bic. Optimum temperature 37 °C. Distinctive characters : Growth very poor unless fresh body fluids are added to the medium. Grows more readily in mixed culture. When grown on a me- dium containing haemoglobin, a black pigment is produced (melanin). Source : Oral cavity, external genitalia, infected surgical wound, urine and feces (Oliver and Wherry). Habitat : Inhabits healthy mucous membranes of mammals, but may take a part in various pathological processes (Burdon). 23. Bacteroides caviae (Vinzent) Hau- duroy et al. (Streptobacillus caviae Vinzent, Ann. Inst. Past., 42, 1928, 533; Hauduroy et al.. Diet. d. Bact. Path., 1937, 53; Spherophorus caviae Pr^vot, Ann. Inst. Past., 60, 1938, 299.) From Cavia, a genus of rodents. Rods: Small, sometimes curved. Usually 0.3 to 0.5 by 1.0 to 1.5 microns. Occurring singly and in chains. Pleo- morphic in old cultures with long filamentous forms. Non-motile. Gram- negative. Serum gelatin : No liquefaction. Serum agar: Surface colonies, small, translucent, slightly raised, adherent to medium in 48 hours. Deep colonies, lenticular, 2 mm in size in 48 hours. Colonies difBcult to break up. No gas. Serum broth : Supernatant fluid clear, with small, stellate colonies, which tend to adhere to walls of the tube. No gas. Indole not formed in serum peptone water. Hydrogen sulfide not formed. Milk: Unchanged. Coagulated egg white and serum not liquefied. No acid or gas from carbohydrates. Pathogenic for guinea pigs, rabbits and mice. Anaerobic. Optimum temperature 37 °C. FAMILY PARVOBACTERIACEAE 575 2. Ristella melaninogenica. See Bac- teroides melaninogenicus. 3. Ristella haloseptica (Wyss) Pr^vot. (Bacterium halosepticum Wyss, Mitt. Grenz. Med. u. Chir., 18, 1904, 199; Prdvot, loc. cit., 291.) From a fatal case of osteomyelitis in man. For a descrip- tion of this species, see Manual, 5th ed., 1939, 570. 4. Ristella putredinis (Weinberg etal.) Pr^vot. (Bacillus A, Heyde, Beitr. z. klin. Chirurg., 76, 1911, 1; Bacillus pu- tredinis Weinberg et al., Les Microbes Ana^robies, 1937, 755; Prevot, loc. cit., 291 . ) Fifteen strains isolated from acute appendicitis. For a description of this species, see Manual, 5th ed., 1939, 571. 5. Ristella terebrans (Brocard and Pham) Prevot. (Bacillus terebrans Bro- card and Pham, Compt. rend. Soc. Biol., Paris, 117, 1934, 997; Prevot, loc. cit., 291.) Two strains isolated from cases of gangrenous erysipelas, associated with a streptococcus. For a description of this species, see IManual, 5th ed., 1939, 571. 6. Ristella furcosa (Veillon and Zuber) Prevot. (Bacillus fiircosus Veillon and Zuber, Arch. M^d. Exp. et Anat. Path., 10, 1898; Fu^iformis furcosus Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 302; Bacter aides furcosus Hauduroy et al., Diet. d. Bact. Path., 1937, 61; Prdvot, loc. cit., 291.) From cases of appendicitis and from lung abscesses. For a description of this species, see Manual, 5th ed., 1939, 572. 7. Ristella putida (Weinberg et al.) Prevot. (Bacillus gracilis putidus Tis- sier and Martelly, Ann. Inst. Past., 16, 1902, 865; Bacillus putidus Weinberg et al., Les Microbes Ana^robies, 1937, 790; not Bacillus putidus Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, * Rearranged by Mrs. Eleanore Heist Clise, New York State Experiment Station, Geneva, New York, December, 1945. Distinctive characters : No growth unless serum is added to the medium. Source : From epidemic benign cervical adenitis of guinea pigs. Habitat : Infected guinea pigs so far as known. Appendix I: Additional species which may belong here. Bacteroides laevis (Distaso) Bergey et al. (Bacillus laevis Distaso, Cent. f. Bakt., I Abt., Orig., 6^, 1912, 444; Bergey et al.. Manual, 1st ed., 1923, 259; not Bacillus laevis -Frankland and Frank- land, Phil. Trans. Roy. Soc. London, 178, B, 1887, 278.) From feces. Bacteroides liquefaciens (Distaso) Ber- gey et al. (Coccobacillus liquefaciens Distaso, Cent. f. Bakt., I Abt., Orig., 59, 1911, 102; Bergey et al.. Manual, 1st ed., 1923, 262.) From feces. Bacteroides rigidus (Distaso) Bergey et al. (Bacillus rigidus Distaso, Cent, f. Bakt., I Abt., Orig., 59, 1911, 103; Bergey et al., Manual, 1st ed., 1923, 263.) Appendix II*: Prevot (Ann. Inst. Past., 60, 1938, 285 and Man. de Class, et de D^term. des Bact. Anaerobies, 1940, 38) has arranged some of the anaerobic, non- spore-forming. Gram -negative, largely parasitic rods in two families, Ristellaceae and Spherophoraceae, as follows : Family Ristellaceae Prevot. (Ann. Inst. Past., 60, 1938, 288.) Genus I. Ristella Prevot. (Lcc. cit., 289.) Straight or slightly bent, non-motile rods. Not capsulated. Gram-negative. Anaerobic. 1. Ristella fragilis. See Bacteroides fragilis. 576 MANUAL OF DETERMINATIVt: BACTERIOLOGY 400; Pr^vot, loc. cit., 291.) From putre- fying meat. For a description of this species, see Manual, 5th ed., 1939, 573. 8. Ristella clostridiiformis (Ankersmit) Prevot. {Bacterium closlndiifonnis Ankersmit, Cent. f. Bakt., I Abt., Orig., 40, 1906, 115; Prevot, Ice. cit., 291.) From the normal intestines of cattle. For a description of this species, see Manual, 5th ed., 1939, 574. 9. Ristella perfoetens (Weinberg et al.) Prevot. {Coccobacillus anaerobius per- foetens Tissier, These MM., Paris, 1900; Bacterium perfoetens Weinberg et al., Les Microbes Ana^robies, 1937, 790; Bacteroides perfoetens Hauduroy et al., Diet. d. Bact. Path., 1937, 67; Prevot, loc. cit., 291.) From the intestines of infants with diarrhoea. For a descrip- tion of this species, see Manual, 5th ed., 1939, 575. 10. Ristella therinophila /3 (Weinberg et al.) Prevot. (Thermo j3, Veillon, Ann. Inst. Past., 36, 1922, 430; Bacillus thermophilus /3 Weinberg et al., Les Microbes Ana^robies, 1937, 800; Prevot, loc. cit., 291.) From manure. Non- pathogenic. 11. Ristella thermophila y (Weinberg et al.) Prdvot. (Thermo y, Veil- lon, Ann. Inst. Past., 36, 1922, 432; Bacillus thermophilus y Weinberg et al., Les Microbes Anat^robies, 1937, 800; Pr(5vot, loc. cit., 291.) From manure. For a description of this species, see Manual, 5th ed., 1939, 575. 12. Ristella incommunis. See Bac- teroides incommunis . 13. Ristella insolita. See Bacteroides insolitus. 14. Ristella halosmophila (Baumgart- ner) Prevot. {Bacteroides halosmophi- lus Baumgartner, Food Research, 2, 1937, 321 ; Prdvot, Man. de Class, et de Determ. des Bact. Anadrobies, 1940, 47.) From salted Mediterranean anchovies. Fre- quently found in the fish muscle and in the solar salt (the probable infecting agent) in which the fish is packed. For a description of this species, see Manual, 5th ed., 1939, 584. 15. Ristella naviformis (Jungano) Prevot. {Bacillus ?iaviformis Jungano, Compt. rend. Soc. Biol., Paris, 66, 1909, 122; Prevot, Ann. Inst. Past., 60, 1938, 291.) From the large intestine of the rat. For a description of this species, see Manual, 5th ed., 1939, 573. 16. Ristella lichenis-plani Prevot. (Bacillus of lichen planus, Jacob and Helmbold, Arch. Derm. Syph., 3, 1933, 28; Prevot, loc. cit., 291.) From the lesions of an inflammatory skin disease, lichen planus. 17. Ristella destillationis (Weinberg et al.) Prevot. (Bacterium, Tunnicliff, Jour. Inf. Dis., 13, 1913, 283; Bacterium destillationis Weinberg et al., Les Microbes Anaerobies, 1937, 762; Prevot, loc. cit., 291.) From a case of chronic bronchitis. 18. Ristella uniformis. See Bacte- roides uniformis. 19. Ristella distasonis. See Bacte- ro ides dis ta,^onis. 20. Ristella uncala. See Bacteroides uncatus. 21. Ristella tumida. See Bacteroides tumidus. 22. Ristella exigua. See Bacteroides exiguus. 23. Ristella trichoides (Potez and Com- pagnon) Prevot. {Bacillus trichoides Potez and Compagnon, Compt. rend. Soc. Biol., Paris, 87, 1922, 339; Bac- FAMILY PARVOBACTERIACEAE 577 teroides trichoides Hauduroy et al., Diet. d. Bact. Path., 1937, 78; Pr^vot, loc.cit., 292.) From a case of choleocysti- tis. For a description of this species, see Manual, 5th ed., 1939, 572. 24. Ristella glutinosa (Guillemot and Hall^) Pr^vot. {Bacillus glutinosus Guillemot and Halle, Arch. M^d. Exp. et Anat. Path., 16, 1904, 599; Bacteroides glutinosus Hauduroy et al.. Diet. d. Bact. Path., 1937, 61; Prevot, loc. cit., 292.) From purulent pleurisies. 25. Ristella capillosa (Tissier) Prevot. (Bacilbis capillosus Tissier, Ann. Inst. Past., £2, 1908, 189; Prevot, loc. cit., 292.) From the intestines of infants. For a description of this species, see Manual, 5th ed., 1939, 573. 26. Ristella cylindroides (Rocchi) Pre- vot. (Bacteriimi cylindroides Rocchi, Lo stato actuale delle nostre cognizioni sui germi anaerobi Gamberine e Parmez- ziani, Bologna, 1908; Prevot, loc. cit., 292.) From the human intestine. For a description of this species, see Manual, 5th ed., 1939, 574, Genus II. Pasteurella Trevisan. Four species. See Bacteroides vul- gatus, Bacteroides ovatus, Bacteroides convexus, and Bacteroides coagulans. Genus III. Dialister Bergey et al. Two species. See Dialister. Genus IV. Capsularis Prevot. (Loc. cit., 290.) Characters as for the genus Ristella, but capsulated. 1. Capsularis zoogleiformans (Wein- berg et al.) Prevot. {Bacillus mucosus anaerobius Prausnitz, Cent. f. Bakt., I Abt., Orig., 89, 1922, 126; Bacterium zoogleijormans Weinberg et al., Les Microbes Anaerobies, 1937, 725; Bac- teroides praussnitzii Hauduroy et al., Diet. d. Bact. Path., 1937, 68; Prevot, loc. cit., 293.) From a purulent em- pyema in man. For a description of this species, see Manual, 5th ed., 1939, 576. 2. Capsularis mucosus (Weinberg et al.) Prevot. {Coccobacterium mucosum anaerobicum Klinger, Cent. f. Bakt., I Abt., Orig., 62, 1912, 186; Bacterium mucosum Weinberg et al., Les Microbes Anaerobies, 1937, 727; Bacteroides vis- cosus Hauduroy et al., Diet. d. Bact. Path., 1937, 81; Prevot, loc. cit., 293.) From a brain abscess following bron- chiectasis in man. For a description of this species, see Manual, 5th ed., 1939, 575. 3. Capsularis variabilis. See Bac- teroides variabilis. Genus V. Zuberella Prevot. {Loc. cit., 290.) Characters as for the genus Ristella, but motile with peritrichous flagella. 1. Zuberella serpens. See Bacteroides serpens. 2. Zuberella praeacuta (Tissier) Pre- vot. {Coccobacillus praeacutus Tissier, Ann. Inst. Past., 22, 1908, 189; Prevot, loc. cit., 293.) From the intestines of infants. For a description of this spe- cies, see Manual, 5th ed., 1939, 577. 3. Zuberella clostridiiformis mobilis Prevot. {Bacterium clostridiiformis Chouk^vitch, Ann. Inst. Past., 25, 1911, 345; Prevot, loc. cit., 293.) From the intestines of a horse. Chouk^vitch con- sidered his organism the same as Anker- smit's Bacterium clostridiiformis, al- though the former was motile. 4. Zuberella aquatilis Prevot. (Spray and Laux, Amer. Water Works Assoc, 578 MANUAL OF DETERMINATIVE BACTERIOLOGY 22. 1930, 235; Pr^vot, loc. cit., 293.) From river water. For a description of this organism, see IVIanual, 5th ed., 1939, 677. 5. Zuberella variegata (Distaso) Pr^- vot. {Bacillus variegatus Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 445; Bacteroides variegatus Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 960; Pr^vot, loc. cit., 293.) From the intestines. For a description of this species, see Manual, 5th ed., 1939, 578. 6. Zuberella rhinitis (Tunnicliff) Pri- vet. (Bacillus rhinitis Tunnicliff, Jour. Inf. Dis., 16, 1915, 493; Pr^vot, loc. cit., 293.) Thirty-two strains isolated from the nasopharynx in human beings suffer- ing from pharyngitis, tonsilitis, bron- chitis and rhinitis, as well as from the nasal mucosa of normal human beings, rabbits, guinea pigs and dogs. For a description of this species, see Manual, 5th ed., 1939, 576. Family Spherophoraceae Pr^vot. (Loc. cit., 289.) Genus I. Spherophorus Prevot. {Loc. cit., 297.) Very pleomorphic rods. Metachro- matic granules common in elongated forms. Non-motile. Non-spore-form- ing. Gram-negative. 1. Spherophorus necrophorus (Fliigge) Prevot. (Bacillus der Kalberdiphthe- rie, Loeffler, Mitteil. kaiserl. Gesund- heitsamte, 2, 1884, 493; Bacillus diphtheriae vitulorum Fltigge, Die Milcro- organismen, 2 Aufl., 1886, 265; Bacillus necrophorus Fliigge, ibid., 273; Bacillus diphtheriae-vitulorum Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Bacillus filiformis Shiitz; not Bacillus filiformis Tils, Ztschr. f. Hyg., 9, 1890, 294; not Bacillus filiformis Migula, Syst. d. Bakt., £, 1900, 587; Nekrosebacillen, Bang, Maanedskrift f . Dyrleager, 2, 1890, 235; Streptothrix cuniculi Schmorl, Deut. Ztschr. f. Tiermed., 17, 1891, 376; Acti- nomyces cuniculi Gasperini, Mittheil. 11 Internal. Med. Congr. Rome, see Cent. f. Bakt., 15, 1894, 684; not Acti- nomyces cuniculi Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 32; Oospora diphtheriae vitulorum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 393; Actinomyces necrophorus Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 434; Streptothrix necro- phora Kitt, Bakterienkunde, 1899; Cory- nebacterium necrophorum Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 531 ; Bacillus necroseos Salomonsen, quoted from Lehmann and Neumann, ibid., 532; Cladothrix cuniculi Mac^, Traite de Bact., 6th ed., 2, 1913, 753; Bacterium necrophorum Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 504 ; Fusiformis necrophorus Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 299; Prevot, Ann. Inst. Past., 60, 1938, 298.) Because of the impor- tance of this organism, a description is included here: Rods: 0.5 to 1.5 microns wide, forming long filaments, up to SO to 100 microns long. Some authors report branching, others deny this. Short forms are re- ported by Schmorl to be motile. Gram- negative. Gelatin stab : No liquefaction. Agar colonies: Small, dirty-white, circular, opaque, with yellowish center under low power lens. Margin floccose. Agar stab : Yellowish colonies along needle track. Gas bubbles produced. Coagulated blood serum: Small, whit- ish colonies, becoming opaque, fimbriate. Broth: Turbid, with gas. Cheese-like odor. Indole is formed. Litmus milk : Cheese-like odor. Acidi- fied and generally coagulated. Nitrites not produced from nitrates. Anaerobic. FAMILY PARVOBACTERIACEAE 579 Optimum temperature 37°C. Produces a soluble exotoxin. Source and habitat : Causes diphtheria in cattle with multiple sclerotic absces- ses; gangrenous dermatitis in horses and mules; multiple necrotic foci in liver of cattle and hogs. One case of human infection reported. Transmissible to mice and rabbits. 2. Spherophorus funduliformis. See Bacteroides funduliformis. 3. Spherophorus necrogenes (Wein- berg et al.) Prevot. (Bacillus, Kawa- mura, Jour. Jap. Soc. Vet. Sci., 5, 1926, 22; Bacillus necrogenes Weinberg et al., Les Microbes Anaerobies, 1937, 381 ; Prdvot, loc. cit., 298.) From epidemic abscesses in hens. 4. Spherophorus necroticus (Nati- velle) Prevot. {Bacillus necroticus Nativelle, 1936, see Weinberg et al., Les Microbes Anaerobies, 1937, 693; Prdvot, loc. cit., 298.) From a case of gangrenous appendicitis. For a description of this species, see Manual, oth ed., 1939, 580. loc. cit., 299.) From a liver abscess in man. For a description of this species, see Manual, 5th ed., 1939, 581. 11. Spherophorus freundi (Hauduroy et al.) Prevot. (Freund, Cent. f. Bakt., I Abt., Orig., 88, 1922, 9; Bacterium of Freund, Weinberg et al., Les Microbes Anaerobies, 1937, 706; Bacteroides freun- dii Hauduroy et al.. Diet. d. Bact. Path., 1937, 57; Prevot, loc. cit., 299.) From a purulent meningitis following otitis in man. For a description of this species, see Manual, 5th ed., 1939, 581. 12. Spherophorus pyogenes (Haudu- roy et al.) Prevot. (Buday, Cent. f. Bakt., I Abt., Orig., 77, 1916, 453; Bacil- lus pyogenes anaerobius Bela-Johan, Cent. f. Bakt., I Abt., Orig., 87, 1922, 290; Bacteroides pyogenes Hauduroy et al., Diet. d. Bact. Path., 1937, 69; Prdvot, loc. cit., 299.) From abscesses of the liver and lungs following septic war wounds. Also from the blood stream following tonsillectomies. For a de- scription of this species, see Manual, 5th ed., 1939, 582. 5. Spherophorus peritonitis Prevot. (Bacillus, Ghon and Sachs, Cent. f. Bakt., I Abt., Orig., 38, 1905, 1 and 131; Prevot, loc. cit., 298.) From peritoneal exudate. 6. Spherophorus gidosus. See Bac- teroides gulosus. 7. Spherophorus inaequalis. See Bac- teroides inaequalis. 13. Spherophorus gonidiaformans (Tunni cliff and Jackson) Prevot. (Ba- cillus gonidiaformans Tunnicliff and Jackson, Jour. Inf. Dis., 36, 1925, 430; Actinomyces gonadiformis (sic) Bergey et al., Manual, 3rd ed., 1930, 469; Bac- teroides gonidiaformans Hauduroy et al., Diet. d. Bact. Path., 1937, 62; Prevot, loc. cit., 299.) From a tonsil. For a description of this species, see Manual, 5th ed., 1939, 582. 8. Spherophorus varius. See Bac- teroides varius. 9. Spherophorus siccus. See Bac- teroides siccus. 10. Spherophorus mortiferus (Harris) Prevot. {Bacillus mortiferus Harris, Jour. Exp. Med., 6, 1901, 519; Prevot, 14. Spherophorus floccosus (Weinberg etal.) Prevot. {Streptobacillus pyogenes floccosus Couromont and Cade, Arch. Med. Exp., 12, 1900, 393; Bacillus floc- cosus Weinberg et al., Les Microbes Anadrobies, 1937, 698; not Bacillus floccosus Kern, Arb. bakt. Inst. Karls- ruhe, 1, Heft 4, 1896, 424; Bacteroides floccosus Hauduroy et al.. Diet. d. Bact. 580 MANUAL OF DETERMINATIVE BACTERIOLOGY Path., 1937, 55; Prevot, loc. cit., 299.) From blood in pyemia of man. For a description of this species, see Manual, 5th ed., 1939, 580. 15. Spherophorus influenzae formis (Russ) Prdvot. {Bacillus influenzae- formis Russ, Cent. f. Bakt., I Abt., Orig., 39, 1905, 357; Bacteroides russii Hauduroy et al.. Diet. d. Bact. Path., 1937, 73; Prevot, loc. cit., 299.) One strain isolated from a perianal abscess and two strains from purulent meningitis in man. For a description of this species, see Manual, 5th ed., 1939, 583. 16. Spherophorus caviae. See Bac- teroides caviae. Genus II. Spherocillus Prevot. (Loc. cit., 297.) Characters as for the genus Sphero- phorus, but motile with peritrichous flagella. 1. Spherocillus bullosus (Distaso) Pre- vot. (Bacillus bullosus Distaso, Cent, f. Bakt., I Abt., Orig., 62, 1912, 443; Bacteroides bullosus Castellani and Chal- mers, Man. Trop. Med., 3rd ed,, 1919, 960; Prevot, loc. cit., 300.) From the intestinal canal. For a description of this species, see Manual, 5th ed., 1939, 583. 2. Spherocillus thetaiotaomicron. See Bacteroides thetaiotaomicron . 3. Spherocillus wirthi Prdvot. (Bacil- lus, Wirth, Cent. f. Bakt., I Abt., Orig., 105, 1928, 201; Prevot, loc. cit., 300.) From a case of acute otitis. Appendix III: The following addi- tional species have been found in the literature. Actinomyces pseudonecrophorus Harris and Brown. (Bull. Johns Hopkins Hosp., 40, 1927, 203.) From the uterus in cases of puerperal infection. Probably should be classified near Spherophorus necrophorus. For a description of this species, see Manual, 5th ed., 1939, 579. Bacillus anaerobius gracilis Lewko- wicz. (Arch. M^d. Exp., 13, 1901, 633.) From the mouths of infants. Bacillus angulosus Garnier and Simon. (Presse Med., 1909, 473.) From the blood of an infant with typhoid fever. Bacillus annuliformans Massini. (Ztschr. f. gesam. Exp. Med., 1913, 81.) From a tuberculous cavity of man. Pathogenic. Bacillus circularis major Heurlin. (Bakt. Unters. d. Keimgehaltes im Genitalkanale d. fiebernden W5chnerin- nen. Helsingfors, 1910, 168.) From the genital canal. Anaerobic. Gram-nega- tive. Bacillus limitans Heurlin. {Ibid., 165.) From the genital canal. Anaero- bic. Gram-negative. Bacillus nebulosus Hall^. (Hall^, These de Paris, 1898; not Bacillus nebu- losus Vincent, Ann. Inst. Past., 21, 1907, 69.) From the human vagina. Bacillus stellatus Vincent. (Vincent, Ann. Inst. Past., 21, 1907, 62; not Bacil- lus stellatus Chester, Man. Determ. Bact., 1901, 274.) From water. Bacillus sijmbiophiles Shottmuller. (Leitfaden f. d. klin. bakt. Kultur., Berlin, 1923.) From the blood in a case of puerperal fever. Bacterium albarrani Jungano. (Compt. rend. Soc. Biol., Paris, 63, 1907, 302.) From a case of cystitis. Bacterium, ininutissimum Le Blaye and Guggenheim. {Cocco-bacillus minu- tissimutn gazogenes Jacobson, Ann. Inst. Past., 22, 1908, 300; Le Blaye and Guggen- heim, Manuel Pratique de Diagnostic Bact., Viget Freres, 1914.) From in- testines of infants. Bacteroides splenomegaliae (Pinoy) Hauduroy et al. {Synbacterium spleno- megaliae Pinoy, Compt. rend. Acad. Sci., Paris, 182, 1926, 1429; Hauduroy et al., FAMILY PARVOBACTERIACEAE 581 Diet. d. Bact. Path., 1937, 75.) From cases of splenomegaly in Algeria. Patho- genic. Pasteurella anaerohiontica Levinthal. (Cent. f. Bakt., I Abt., Orig., 106, 1928, 195.) From the naso-pharj^nx of man. Streptobacillus gracilis Guillemot and Hall^. (Guillemot and Hall4, Arch. Med. Exp. et Anat. Path., 16, 1904, 598; Bacteroides streptobacilliformis Haudu- roy etal., Diet. d. Bact. Path., 1937, 76.) From putrid pleurisies. Genus II. Fusobacterium Knorr.* (Knorr, Cent. f. Bakt., I Abt., Grig., 87, 1922, 53Q; Fusiformis Prevotand Fusocillus Pr(5vot, Ann. Inst. Past., 60, 1938, 300.) Gram-negative, anaerobic rods, usually with tapering ends. Usually non-motile. Stain with more or less distinct granules. The type species is Fusobacterium plauti-vincenti Knorr. Key to the species of genus Fusobacterixun. I. Acid from maltose. A. No gas produced. 1. Fusobacterium plauti-vincenti. B. Gas produced. 2. Fusobacterium biacutum. II. No acid from maltose. A. Disagreeable odor produced on cultivation. 3. Fusobacterium nucleatum. B. No odor produced. 4. Fusobacterium polymorphum. 1. Fusobacterium plauti-vincenti Knorr. (Knorr, Cent. f. Bakt., I Abt., Orig., 89, 1923, 16; Fusiformis plauti- vincenti and Fusiformis vincenti Haudu- roy etal., Diet. d. Bact. Path., 1937, 240.) Named for H. C. Plaut and for H. Vincent who studied diseases of the respiratory tract. The relationships between this organ- ism and the following have not been clearly established: Fusiformis dentium Hoelling, Arch. f. Protistenkunde, 19, 1910, 240; Bacillus fusiformis Veillon and Zuber, Arch, de med. e.xper., 10, 1898, 517 {Corynebaclerium fusiforme Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 529); not Bacillus fusi- formis Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 724; Fusiformis fusiformis Topley and Wilson, Princip. of Bact. and Immun., 1st ed., 1, 1931, 300. Weinberg, Nativelle and Pr^vot (Les Microbes Ana^robies, 1937, 804) and Prevot (Ann. Inst. Past., 60, 1938, 285) make a distinction between Plaut *s bacillus {Fusocillus plauti) and Vin- cent's bacillus (Fusiformis fusiformis), the former being actively motile and non- pathogenic and the latter non-motile and pathogenic. Rods: 0.5 to 1.0 by 8 to 16 microns, occurring in pairs with blunt ends to- gether and outer ends pointed, sometimes in short, curved chains or long spirillum- like threads. Granules present. Non- motile. Gram-negative. Serum agar shake culture : After 36 hours, colonies spherical, up to 0.5 mm in diameter, thin, j-ellowish-brown. Serum agar plate : Matted growth. Medium around colonies becomes turbid from the precipitation of protein. No surface growth. Serum broth : Milkj^ turbidity. Liver broth: No turbidity. Grayish- white, flakj^ precipitate. * Arranged by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York, December, 1938; rearranged, December, 1945. 582 MANUAL OF DETERMINATIVE BACTERIOLOGY Indole not formed. Acid from glucose, fructose, sucrose, maltose and sometimes from lactose. No acid from inulin or mannitol. (Hine and Berry, Jour. Bact., 34, 1937, 524.) No H2S produced. No odor produced. No gas formed. Non-pathogenic for white mice (Hau- duroy et al., loc. cit.). Temperature relations: Optimum 35° to 37°C. Minimum 30°C. (Hauduroy et al., loc. cit.) Optimum pH 6.8 to 8.0 (Hauduroy et al., loc. cit.). Anaerobe . Source : Two strains isolated from deposit on teeth. Habitat : Presumably the buccal cavity. 2. Fusobacterium biacutum Weinberg and Prdvot. (Weinberg and Prdvot, Compt. rend. Soc. Biol., Paris, 95, 1926, 519 ; Fusiformis biacutus Hauduroy et al.. Diet. d. Bact. Path., 1937, 238.) From Latin, double pointed. Rods: 0.4 to 0.5 by 1.4 to 3.0 microns, with pointed ends, occurring singly, in pairs or sometimes in short chains. Non- motile. Gram-negative. Gelatin : No liquefaction. Veillon's agar: Rapid growth. Colo- nies lens-shaped. Gas is produced which breaks up the medium. Plain broth: Poor growth. Glucose broth: Turbid. Medium rapidly acidified. Good growth. Indole not formed. Milk: Acid and coagulation in 2 to 8 days. Curd not digested. Casein and coagulated egg-white not digested. Neutral red reduced. Acid from glucose, fructose, galactose, maltose and lactose. Small amount of H2S produced. Does not require blood serum for growth. Pathogenic for guinea pigs. Killed in 60 minutes at 60°C. Anaerobic. Source : Six strains isolated from a case of appendicitis. Habitat : Unknown. 3. Fusobacteriiim nucleatum Knorr. (Knorr, Cent. f. Bakt., I Abt., Orig., 89, 1923, 17; Fusiformis nucleatus Bergey et al.. Manual, 3rd ed., 1930, 514.) Lat- inized, nucleated. Rods: 1.0 by 4.0 microns, spindle- shaped, occurring singly. One or two granules present. Non-motile. Gram- negative. Serum agar plate : Deep colonies lens- shaped with offshoots. Plain liver broth: No growth. Liver broth with serum: After 1 to 3 days, flocculent deposit on the pieces of liver. Indole not formed (Knorr, loc. cit.). Indole formed (Hine and Berry, Jour. Bact., 34, 1937, 521). • Disagreeable odor produced on cul- tivation. No gas produced. Acid from glucose, usually from fruc- tose, sometimes from sucrose and lactose. No acid from maltose, inulin or mannitol. (Hine and Berry, loc. cit.) No H2S formed. Temperature relations: Optimum 35° to 37 °C. Minimum 30°C. (Hauduroy et al., Diet. d. Bact. Path., 1937, 239.) Survives 56°C for 15 minutes, but not 60°C for 10 minutes (Hine and Berry, loc. cit.). Optimum pH 6.8 to 8.2 (Hauduroy et al., loc. cit.). Anaerobe. Source : One strain isolated from de- posit on teeth in a healthy mouth. Habitat : Presumably the buccal cavity. 4. Fusobacterium polymorphum Knorr. (Knorr, Cent. f. Bakt., I Abt., Orig., 89, 1923, 19; Fusiformis polymorphus Bergey et al.. Manual, 3rd ed., 1930, 515.) From Greek, assuming many forms. Rods: 0.2 to 0.5 by 8 to 16 microns. FAMILY PARVOBACTERIACEAE 583 occurring in pairs with the pointed ends adjoining. Often occurring in threads. Non-motile. Gram-negative. Serum agar plates (alkaline) : After 2 to 3 days, colonies 0.5 mm or larger, lens- shaped with offshoots. Tenacious sediment in liquid media. Indole not formed (Knorr, loc. cit.). Indole formed (Hine and Berry, Jour. Bact., Sit, 1937, 522). No gas produced. No odor produced. Acid usually produced from glucose, fructose, and sucrose. No acid from lactose, maltose, inulin or mannitol. (Hine and Berry, loc. cit.) No H2S produced. Temperature relations: Optimum 35° to 37°C. Minimum 30°C. (Hauduroy et al., Diet. d. Bact. Path., 1937, 242.) Survives SO^C for 15 minutes, 52°C for 10 minutes and 56°C for 5 minutes (Hine and Berry, Zoc. a'f., 523). Optimum pH 7.0 to 8.2 (Hauduroy et al., loc. cit.). Anaerobe. Source : One strain isolated from de- posit on teeth in a case of gingivitis. Habitat : Presumably the buccal cavity. Appendix I: The following species are mentioned here because they appear to be related to the organisms in the genus Fusobacterium : Bacillus hastilis Seitz. (Ztschr. f. Hyg., 30, 1899, 47; Mycohacleriwn hastilis Chester, jVIan. Determ. Bact., 1901, 356.) A collective name for the organisms frequently found in stinking pus and in tonsillar pockets in both healthy and diseased mouths. Fusocillus shmamini Prevot. (Ann. Inst. Past., 60, 1938, 300.) Feebly motile. Fusiformis acnes, Fusiforrnis hodgkini and Fusiformis typhi-exanthematici (Plotz) of Holland (Jour. Bact., 5, 1920, 223) are names presumably intended for bacteria more properly placed in the genus Corynebacterium. Fusiformis muris Hoelling. (Arch. f. Protistenkunde, 19, 1910, 239.) From the blind gut of a mouse. Stated by the author to be similar to Fusiformis termi- tidis Hoelling. Fusiformis necrophorus Topley and Wilson. See Sphcrophorus necrophorus Prevot, page 578. Fusiformis nodosus Beveridge. (Bev- eridge, Austral. Council Sci. and Indus. Res. Bui. 140, 1941, 56 pp.; Actinomyces nodosus Hagan, The Infectious Dis- eases of Domestic Animals. Ithaca, New York, 1943, 312.) Considered the primarj' cause of footrot of sheep. Also see Spirochaeta penortha. Appendix II: Because of the preferable form of the name and also because it is questionable whether the anaerobic fusi- form bacteria of the mouth closely re- semble the more or less aerobic bacteria found in termites, the genus name Fuso- hacterium Knorr has been used for the mouth organisms. The termite organ- isms live in the intestinal tract bathed in digested wood and have the micro- scopic appearance of the cellulose- destroying Cytophaga Winogradsky. These are shown by Stanier (Jour. Bact., 40, 1940, 619) to belong to Myxobacte- riales. The organisms placed in Fusiformis Hoelling are as follows: Fusiformis hilli Duboscq and Grasse. (Arch. Zool. Exper. et Gen., 66, 1927, 454 and 486.) Found as an ectoparasite on flagellates Descovina spp. which live in the rectum of termites {Calotermes ( Glyptotermes ) iridipennis ) . Fusiformis iermitidis Hoelling. (Arch, f. Protistenkunde, 19, 1910, 239.) From the intestinal tract of termites. 584 MANUAL OF DETERMINATIVE BACTERIOLOGY TRIBE IV. HEMOPHILEAE WINSLOW ET AL. (Jour. Bact., 5, 1920, 212.) Minute parasitic forms growing on first isolation only in the presence of hemoglobin, ascitic fluid or other body fluids, or in the presence of certain growth accessory sub- stances found in sterile, unheated plant tissue (potato) . Motile or non-motile. Com- monly found in the mucosa of respiratory tract or conjunctiva. Key to the genera of tribe Hemophileae. I. Aerobes to facultative anaerobes. A. Non -motile. 1. Predominantly occurring singly. Genus I. Hemophilus, p. 584. 2. Predominantly occurring as diplobacilli. Genus II. Moraxella, p. 590. B. Motile, encapsulated. Genus III. Noguchia, p. 592. II. Anaerobes. A. Non -motile. Genus IV. Dialister, p. 594. Genus I. Hemophilus Winsloio et al.* (Jour. Bact., 2, 1917, 561.) From Greek, loving blood. Minute rod-shaped cells, sometimes thread-forming and pleomorphic. Non-motile. Gram-negative. Strict parasites growing best (or only) in the presence of hemo- globin and in general requiring blood serum, ascitic fluid, or certain growth accessory substances. The type species is Hemophilus influenzae (Lehmann and Neumann) Winslow et al. Key to the species of genus Hemophilus. I. Affecting the respiratory tract. 1. Require both V and X growth factors for growth. 1. Hemophilus influenzae. 2. Hemophilus suis. 3. Hemophilus hemolyticus. 2. V growth factor sufficient for growth. 4. Hemophilus parainfluenzae. 5. Hemophilus pertussis. II. Affecting the genital region. 3. X growth factor sufficient for growth. 6. Hemophilus ducreyi. 7. Hemophilus haemoglobinophilus. * Revised by Dr. Margaret Pittman, National Institute of Health, Bethesda, Maryland, October, 1945. FAMILY PARVOBACTERIACEAE 585 Where the relationship to growth accessory factors is known, the following table may serve as a key : Growth in peptone water containing Species Growth (actor X Phospho- pyridine nu- cleotide (V) Growth factors XandV Hemophilus influenzae + 1 1 1 1 ± + + Hemophilus suis + Hemophilus hemolyticus . . + Hemophilus parainfluenzae + Hemophilus haemoglobinophilus + 1. Hemophilus influenzae (Lehmann and Neumann) Winslow et al. (Koch, Wiener med. Wchnschr., 33, 1883, 1550, Weeks, New York Med. Record, 31, 1887, 571; Influenzabacillus, Pfeiffer, Deutsche med. Wchnschr., 1892, 28; Ztschr. f. Hyg., 13, 1893, 357; Bacterium influenzae Lehmann and Neumann, Bnkt. Diag., 1 Aufl., 2, 1896, 187; Bacillus in- fluenzae Kruse, in Flligge, Die Mikro- organismen, 3 Aufl., 2, 1896, 434; Bac- terium aegyptiacum Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 191 ; Hemophilus meningitidis cerebro- spinalis septicemiae Cohen, Ann. Inst. Past., 23, 1909, 273; Winslow et al., Jour. Bact., 2, 1917, 561; Coccobacillus pfeifferi Neveu-Lemaire, Precis Parasi- tol. Hum., 5th ed., 1921, 20; Hemophilus conjunctivitidis Bergey et al.. Manual, 1st ed., 1923, 270.) From influenza, a disease of the respiratory tract. Common name : The Koch-Weeks Bacillus. Very small rods: 0.2 to 0.3 by 0.5 to 2.0 microns, occurring singly and in pairs, occasionally in short chains, and at times long thread forms are seen. Frequently show a marked tendency to bipolar stain- ing. Some strains are encapsulated. Non-motile. Gram-negative. Requires both the factors X and V for its growth. Gelatin colonies: No growth. Gelatin stab : No growth . Blood agar colonies: Small, circular, transi)arent, homogeneous, entire. Satellitism with Staphylococcus. Blood agar slant : Thin, filiform, trans- parent growth. Chocolate agar slant : Luxuriant growth. Blood broth: Slightly turbid. No hemolysis. litmus milk, with blood : Some strains render it very slightly alkaline. Sterilized potato slant : No growth. Fresh unheated sterile potato added to broth favors development. Indole is formed by some strains. Nitrites are produced from nitrates. Some strains attack none of the carbo- hydrates, while other strains attack various carbohydrates, provided a suit- able medium is used. IVIannitol and lactose never fermented. Pathogenic. Aerobic, facultative. Optimum temperature 37°C. Maxi- mum 43°C. Minimum 26° to 27°C. Killed at 55°C for thirty minutes. Source : Isolated by Pfeiffer in cases of influenza. Found in conjunctiva, naso- pharynx, sputum, sinuses, cerebrospinal fluid, blood, and pus from joints. Habitat: Respiratory tract. A cause of acute respiratory infections, of acute conjunctivitis, and of purulent meningi- tis of children, rarely of adults. Re- 586 MANUAL OF DETERMINATIVE BACTERIOLOGY garded by Pfeiffer und others to be the cause of influenza. Note: Six types (a-f) of Hemo-philus influenzae are recognized on the basis of precipitation of immune serum by cap- sular substance. Strains from cerebro- spinal fluid are usually of type b. The majority of the strains from the icspira- tory tract are not type-speciflc. 2. Hemophilus suis Hauduroy et al. {Hemophilus influenzae suis Lewis and Shope, Jour. Exp. Med., 54, 1931, 361 and 373; Bacterium influenzae suis Kobe, Cent. f. Bakt., I Abt., Orig., 129, 1933, 161 ; Hauduroy et al., Diet. d. Bact. Path., 1937, 258.) From Latin, swine. Resembles Hemophilus influenzae ex- cept it is relatively inert to growth, in- dole is not formed, nitrites are produced from nitrates, and maltose and sucrose are slightly fermented but not the car- bohydrates fermented by Hemophilus influenzae. Source: From cases of swine influenza. Habitat: With a filterable virus it causes swine influenza. 3. Hemophilus hemolyticus Bergey et al. (Bacillus X, Pritchett and Still- man, Jour. Exp. Med., 29, 1919, 259; Stillman and Bourn, Jour. Exp. Med., 32, 1920, 665; Bergey et al.. Manual, 1st ed., 1923, 269.) From Greek, blood-dis- solving. Morphologically like Hemophilus in- fluenzae. Non-motile. Gram-negative. Requires both the factors X and V for its growth. Valentine and Rivers (Jour. Exp. Med., 45, 1927, 993) isolated certain hemolytic strains which did not entirely agree with this. Blood agar colonies : Resemble Hemo- philus influenzae but surrounded by a zone of hemolysis. Blood agar slant : Thin, filiform, trans- parent growth. Blood broth: Turbid, showing hemol- ysis. Blood milk mixture : Slightly alkaline. Sterile luiheated potato favors de- velopment . Indole is formed by some strains. Nitrites are produced from nitrates. Some strains do not attack carbohy- drates, other strains ferment various carbohydrates. Aerobic, facultative. Optimum temperature 37 °C. Habitat : Found in upper respiratory tract of man. No n -pathogenic. 4. Hemophilus parainfluenzae Rivers. (Johns Hopkins Hosp. Bull., 33, 1922, 429.) From Latin, like, and the disease, influenza. Morphologically like Hemophilus in- fluenza. Non-motile. Gram-negative. Requires the V factor for its growth. Blood agar colonies : Resemble Hemo- philus influenzae. No hemolysis. Blood agar slant : Thin filiform trans- plant. Broth containing yeast extract : Floc- cular sediment. Indole is formed by some strains from cat. Nitrites are produced from nitrates. Some strains attack none of the carbo- hydrates; other strains ferment various carbohydrates . Aerobic, facultative. Optimum temperature 37°C. Habitat : Found in upper respiratory tract of man and cat. Usually non- pathogenic. 5. Hemophilus pertussis Holland. (Microbe de coqueluche, Bordet and Gengou, Ann. Inst. Past., 20, 1906, 731; Holland, Jour. Bact., 5, 1920, 219; Bacillus pertussis Holland, idem; Bac- terium tussis convulsivae Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 317 (Bordet and Gengou's organism) ; not Bacterium tussis convulsivae Leh- mann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 192 (Czaplewski and FAMILY PARVOBACTERIACEAE 587 Hensel's organism).) From Latin, whooping cough. Short, oval rods: 0.2 to 0.3 by 1.0 mi- cron, occurring singlj^ or sometimes in pairs and short chains. Show tendency to bipolar staining. Capsules may be demonstrated by special technic (Law- son). Non-motile. Gram -negative. No growth on the usual laboratorj- media; adapted by repeated transfer with heavy inoculum. Adaptation accompa- nied by loss of original characteristics. Bordet-Gengou medium or some modi- fication containing at least 15 per cent blood is optimum for isolation and main- tenance. Colonies: Smooth, raised, entire, pearly, glistening. Surrounded by a zone of hemolysis. Litmus milk: (After adaptation) brownish color. Alkaline. Indole not formed. Nitrites not produced from nitrates. No action in carbohydrate media. Catalase positive. Aerobic. Optimum temperature 37 °C. Serologically homogeneous when first isolated (Phase I of Leslie and Gardner). Dissociative changes, recognizable mor- phologically, culturally, antigenically, and by animal tests, take place when maintained on unfavorable media. Source : From respiratory tract in cases of whooping cough, especially bj- the cough plate method. Habitat : Etiologically associated with whooping cough. Note: Bacillus para-pertussis Elder- ing and Kendrick. (Jour. Bact., 35, 1938, 561.) From cases of whooping- cough. Closely related antigenically to Hemophilus pertussis and Brucella bron- chiseptica, but distinct from either. 6. Hemophilus ducreyi (Neveu- Lemaire) Bergey et al. (Ducrey, Cong, internat. de dermatol. et syph., Compt. rend., Paris, 1890, 229; Monatshft. f. prakt. Dermatol., 9, 1889, 387; Riforma med., 5, 1889, 98; Monatshft. f. prakt. Dermatol., 21, No. 2; Streptobacillus of soft chancre, Ducrey, Abstract in Cent. f. Bakt., I Abt., 18, 1895, 290; Bacillus idceris cancrosi Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 456; Bacterium ulceris cancrosi Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 67; Bacterium cancrosi Chester, Manual Determ. Bact., 1901, 120; Coccobacillus ducreyi Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 20; Bergey et al., Manual, 1st ed., 1923, 271.) Named for Ducre}^ who first isolated the organism. Small rods: 0.5 by 1.5 to 2.0 microns, with rounded ends, occurring singly and in short chains. Non-motile. Gram- negative. Requires the X factor for its growth. Gelatin colonies: No growth. Gelatin stab: No growth. Blood agar colonies: Small, grayish, glistening, showing a slight zone of hemol- ysis around the colony in three or four days. Best growth is obtained on clotted rab- bit, sheep, or human blood heated to 55°C for 15 minutes, and in casein digest agar containing blood. Moisture is im- portant for growth. Aerobic, facultative. Optimum temperature 37°C. Habitat : The cause of soft chancre (chancroid). 7. Hemophilus haemoglobinophilus (Lehmann and Neumann) Murray. {Bacillus haemuglobinophilus canis Friedberger, Cent. f. Bakt., I Abt., Orig., 33, 1903, 401; Bacterium haemoglo- binophilus Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 270; Hemophilus canis Rivers, Johns Hopkins Bull., 33, 1922, 149; Jour. Bact., 7, 1922, 579; Mur- ray, in Manual, 6th ed., 1939, 309.) From Greek, hemoglobin-loving. Small rods: 0.2 to 0.3 by 0.5 to 2.0 microns, occurring singly, in pairs and 588 MANUAL OF DETERMINATIVE BACTERIOLOGY short chains. Non-motile. Gram- negative. Requires the X factor for its growth. Blood agar colonies : Small, clear, trans- parent, entire. Old colonies become opaque. Blood broth : Turbid. Blood milk mixture: Doubtful de- velopment. Indole is formed. Nitrites produced from nitrates. Acid but no gas from glucose, fructose, galactose, mannitol, sucrose and xylose. No acid from maltose, lactose dextrin, arabinose or glycerol. (Rivers, Zoc. cit.) Optimum temperature 37°C. Aerobic, facultative. Habitat : Occurs in large numbers in preputial secretions of dogs. Appendix I:* The following species has been placed in the tribe Hemophileae by Van Rooyen (Jour. Path, and Bact., 43, 1936, 469). It has been pointed out by Buchanan (General Systematic Bac- teriology, 1925, 490) that the genus name Streptobadllus is invalid. Streptobacillus moniliformis Levaditi, Nicolau and Poincloux. (Compt. rend. Acad. Sci., Paris, 180, 1925, 1188.) This organism is regarded as identical with Haverhillia multiformis Parker and Hudson (Amer. Jour. Path., 2, 1926, 357) by Van Rooyen {loc. cit.). Topley and Wilson (Princip. Bact. and Immun., 2nd ed., 1936, 270) regard it as identical with their Actinomyces muris {Strepto- thrix ratti Schottmtiller), the cause of rat-bite fever. Asterococcus muris Heil- man. Jour. Inf. Dis., 69, 1941, 32. See Actinomyces muris ratti in the Appendix to the genus Streptomyces. Jordan and Burrows (Textb. of Bact., 14th ed., 1946, 614) consider all these names synony- mous. Dawson and Hobby (Proceed- ings, Third Internal. Congr. for Micro- biol., New York, 1940, Section I, 177) suggest that the pleuropneumonia-like cultures isolated from Streptobacillus moniliformis really represent variant phases in the growth of this organism. Description from Levaditi et al. {loc. cit.) and Brown and Nunemaker (Bull. Johns Hopkins Hosp., 70, 1942, 201). Streptobacilli : 2.0 to 3.0 microns in length, pleomorphic, with branching filaments up to 30 to 40 microns long, fragmented, bacillary and coccobacillary forms. Swollen and club-shaped cells are found. Morphology is best demon- strated by aniline dyes, e.g. Wayson's plague stain. Non-motile. Gram-nega- tive. Enriched media are required for good growth. Best liquid media are rabbit blood and broth containing serum or ascitic fluid. Best solid media are glycerol extract of potato-infusion broth-egg yolk medium and nutrient agar containing serum. Blood agar or ascitic serum agar : Col- onies small, clear. Blood plates: Growth slow. Numer- ous small whitish colonies appear on the ■ third day. Veillon's medium: Punctiform col- onies, abundant in depth, less growth at surface. No gas. Broth with ascitic fluid and globular extract : Good growth, forming clots which precipitate and are rather adherent to one another. Growth rapidly re- duces the pH of the medium killing the bacteria in cultures 24 hrs. old. Milk: Slow growth. No coagulation. Loffler's serum: Poor growth. Virulent for rabbits and mice. Good growth at 37 °C. Facultative anaerobe. Grows better under anaerobic conditions in the pres- ence of added CO2, than in the presence of air. Source : Isolated from a case of a febrile, * Appendixes I and II arranged by Prof. E. G. D. Murray, McGill Univ., Montreal, P. Q., Canada, March, 1946. FAMILY PAKVOBACTERIACEAE 589 septicemic disease, accompanied by ar- thritis, erythema and angina. Habitat : The cause of an acute febrile disease sometimes called erythema mul- tiforme. Appendix II: The following species may be identical with some of those listed above or related to them : Bacillus marianensis Leber and Pro- wazek. (Berlin, klin. Wochnschr., 1, 1911, 27.) Allied to the Koch-Weeks Bacillus. Associated with cases of pink eye. Bacillus weeksi Xeveu-Lemaire. (Precis Parasitol. Hum., 5th ed., 1921, 24.) Described bj' Weeks. The cause of trachoma or granular conjunctivitis in tropical countries. Transmitted by the domestic fly. Probably intended, for the Week's bacillus (Weeks, New York Med. Record, 31, 1887, 571). Bacterium tiissis convulsivae Lehmann and Neumann. (Bacterium, Czaplewski and Hensel, Deut. med. Wochnschr., 23, 1897, 587; Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 192; not Bacterium tussis convulsivae Lehmann and Neumann, ibid., 7 Aufl., 2, 1927, 317; Bacillus tussis convulsivae Leh- mann and Neumann, ibid., 4 Aufl., 2, 1907, 269.) Considered the cause of whooping cough by the original isolators. Hemophilus aphrophilus Khairat. (Jour. Path, and Bact., 50, 1940, 497.) From blood and from heart valve of a case of endocarditis. Hemophilus cuniculi Hauduroy et al. {Hemophilus sp. Gibbons, Jour. Inf. Dis., 45, 1929, 288; Hauduroy et al.. Diet. d. Bact. Path., 1937, 249.) From skin abscesses of rabbits. Hemophilus gallinarum Delaplane, Er- win and Stuart and Hemophilus galli- narum Eliot and Lewis. (Bacillus hemo- globinophilus coryzae gallinarum De Blieck, Tijdsch. v. Diergeneensk., 58, 1931, 310; also see Vet. Jour., 88, 1932, 9; Delaplane, Erwin and Stuart, R. I. State Coll. Sta. Bull. 244, May, 1934; Eliot and Lewis, Jour. Amer. Vet. Med. Assoc, 84, 1934, 878.) From edematous fluid from the head of a chicken. The cause of an infectious rhinitis in chickens. Hemophilus influenzae murium (Ivairies and Schwartzer) LwofT. {Bac- terium influenzae murium Kairies and Schwartzer, Cent. f. Bakt., I Abt., Orig., 137, 1936, 351; Lwoff, Ann. Inst. Past., 62, 1939, 168.) From the lungof a mouse. Hemophilus meningitidis (Martins) Hauduroy et al. {Coccobacillus menin- gitidis Martins, Compt. rend. Soc. Biol., Paris, 99, 1928, 955; Hauduroy et al., Diet. d. Bact. Path., 1937, 254.) Re- sembles Hemophilus influenzae except that it shows sluggish motility. From ce re bro -spinal fluid. Hemophilus muris Hauduroy et al. (Bacillus of an epizootic of stock mice, Mackie, Van Rooyne and Gilroy, Brit. Jour. E.xp. Path., I4, 1933, 132; Haudu- roy et al., Diet. d. Bact. Path., 1937, 255. ) From heart blood, spleenand other organs of mice dying from an epizootic disease . Hemophilus ovis Mitchell. (Jour. Amer. Vet. Assoc, 68, 1925, 8.) From bronchi of sheep. Hcjnophilus pertussis Ford. {Bacillus pertussis eppendorf Jochmann and Krause, Ztschr. f. Hyg., 36, 1901, 193; Ford, Textb. of Bact., 1927, 615; not necessarily identical with Bordet and Gengou's organism, Hemophilus pertussis Holland, Jour. Bact., 5, 1920, 215.) From the respiratory tract and lungs in per- tussis. Hemophilus putoriorum Hauduroy et al. {Bacterium influenzae putoriorum multiforme Kairies, Ztschr. f. Hyg., 117, 1935, 12; Hauduroy et al., Diet. d. Bact. Path., 1937, 258.) From the respiratory tract of ferrets. Lehmann and Neumann (Bakt. Diag., 6 Aufl., 2, 1920, and 7 Aufl., 2, 1927) list 590 MANUAL OF DETERMINATIVE BACTERIOLOGY the following species as closely related to this group : Bacillus caiarrhalis JundeW . (Hygieae, 60, No. 6 and 7, p. 667.) From cases of acute bronchitis. Bacillus trachomatis Lehmann and Neumann. (The Bacillus Miiller, Luers- sen, Cent. f. Bakt., I Abt., Orig., 39, 1905, 682.) From conjunctiva. Bacterium czaplewskii Chester. (Ba- cillus bei Keuchhusten, Czaplewski, Cent. f. Bakt., ^2, 1897, 641; Bacterium tussis convulsivae Lehmann and Neu- mann, Bact. Diag., 2 Aufl., 1899, 192; Chester, Man. Determ. Bact., 1901, 153.) From sputum in whooping cough. This is not now regarded as being etiolog- ically associated with whoopitig cough. Bacterium exiguum Staubli. (Miinch. med. Wchnschr., No. 45, 1905.) From a case of septic endocarditis. Bacterium microbutyricum Hellstein. From butter. Bacterium minutissimus sputi (Luz- zatto) Lehmann and Neumann. {Ba- cillus minutissimus sputi Luzzatto, Cent, f. Bakt., I Abt., 37, 1900, 816.) From a case of pertussis. Bacterium polymorphwn convulsivum Melfi. (Cent. f. d. gesamte Hygiene, 7, 1924, 133.) Bacterium septicaemiae canis Paranhos . (Cent. f. Bakt., I Abt., Orig., 50, 1909, 607.) Streptobacillus ureihrae Pfeiffer. (Cent. f. Bakt., I Abt., Ref., S6, 1905, 59.) From the normal urethra and from cases of chronic cystitis and urethritis. Genus II. Moraxella Lwoff* {Diplobacillus McNab, Klinische Monatsbl. f. Augenheilk., 4^, 1904, 64; not Diplo- bacillus Weichselbamn, Cent. f. Bakt., 2, 1887, 212; Lwoff, Ann. Inst. Past., 62, 1939, 168.) Named for Morax, who first isolated the type species. Small, short, rod-shaped cells, usually occurring singly or in pairs. Non-motile. Parasitic. Aerobic. Gram-negative. The type species is Moraxella lacunata (Eyre) Lwoff. Key to the species of genus Moraxella. I. No growth in gelatin. 1. Moraxella lacunata. II. Gelatin liquefied. A. Rapid liquefaction. No growth in milk. 2. Moraxella liquefaciens . B. Very slow liquefaction. Cells capsulated. Growth in milk. 3. Moraxella bovis. 1. Moraxella lacunata (Eyre) Lwoff. (Diplobacille de la conjunctivite sub- aigue, Morax, Ann. Inst. Past., 10, 1896, 337; Diplobacillus of chronic conjunctivi- tis, Axenfeld, Cent. f. Bakt., I Abt., 21, 1897, 1 ; Bacterium conjunctivitis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Bacillus lacunatus Eyre, Jour. Path, and Bact., 6, 1899, 1; not Bacillus lacunatus Wright, Memoirs Nat. Acad. Sci., 7, 1895, 435; Diplobacillus morax- axenfeld McNab, Klinische Monatsbl. f. Augenheilk., 4^, 1904, 64; Bacterium duplex Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 193; Hemophilus lacunatus Holland, Jour. Bact., 5, 1920, 223; Bacillus duplex Hewlett, Med. Res. Council Syst. of Bact., 2, 1929, 417; * Arranged by Prof. E. G. D. Murray, McGill University, Montreal, P. Q., Canada, September, 1945. FAMILY PARVOBACTERIACEAE 591 Lwoff, Ann. Inst. Past., 62, 1939, 173; Hemophilus duplex Murray, in Manual, 5th ed., 1939, 308.) From Latin, pitted. Audureau (Ann. Inst. Past., 64, 1940, 128) recognizes an atypical variety of this species. To distinguish between the two varieties, she designates these as Moraxella lacunata var. typica and Moraxella lacunata var. atypica. Short rods: 0.4 to 0.5 by 2.0 microns, occurring singly and in pairs and short chains. Ends rounded or square in the chains. Non-motile. Gram-negative. Gelatin colonies: No growth. Gelatin stab: No growth. Blood agar colonies: Small, circular, transparent, entire. Growth on sub- culture difficult. Certain strains are not surrounded by zones of hemolj^sis; others are (Oag, Jour. Path, and Bact., 54, 1942, 128). Serum agar colonies: Delicate, grayish. Loffler's blood serum: Slow but definite liquefaction (pitting) around the colonies. Ascitic broth : Turbid with slight, grayish sediment. Blood milk mixture : Doubtful de- velopment. Litmus milk: Unchanged. Potato: No growth. Indole not formed. Nitrites not produced from nitrates. Various carbohydrates and mannitol are attacked. Optimum temperature 37 °C. Aerobic, facultative. Source : From conjunctiva. Habitat : The cause of subacute in- fectious conjunctivitis, or angular con- junctivitis. 2. Moraxella liquefaciens (McNab) comb. nov. (Diplobacille liqu^fiant. Pet- tit, Annales d'oculistique, March, 1899, 166 and Thesis, Paris, 1900, 223; Diplo- bacillus liquefaciens McNab, Klinische Monatsbl. f. Augenheilk., 42, 1904, 64; Bacillus duplex liquefaciens Lwoff, Ann. Inst. Past., 62, 1939, 170; Moraxella duplex liquefaciens Lwoff, ibid., 171; Moraxella duplex Lwoff, ibid., 171; Moraxella duplex var. liquefaciens Audu- reau, Ann. Inst. Past., 64, 1940, 139.) From Latin, liquefying. Diplobacilli : 1.0 to 1.5 by 2.0 microns, occurring singly and in pairs, and having rounded ends. Capsules not demon- strated. Non-motile. Stain uniformly with basic aniline dyes. Gram-negative. Gelatin colonies: Round, 1.5 to 2.0 mm in diameter, yellowish-white. Gelatin stab: Rapid liquefaction. Blood agar: Ready growth in primary and subculture. Ascitic agar colonies : Grayish, thick, round, viscous. Peptone agar colonies : Same as above, but less abundant growth. Coagulated serum: Liquefaction in 3 to 4 daj's; eventually complete. Plain broth : Poor growth, if any. Slight uniform turbidity. Ascitic broth : Abunda,nt growth in 24 hours at 35°C. Uniform turbidity. Later sediment and an opaque pellicle. Milk: No growth. No coagulation. Potato : Slight, yellowish-white, vis- cous growth. Optimum temperature between 20° and 37°C. ffilled at 55°C for 15 minutes. Aerobic . Not pathogenic for laboratory animals. Source: From cases of conjunctivitis associated with corneal ulceration in man. Habitat : Conjunctivitis in man so far as known. 3. Moraxella bovis (Hauduroy et al.) comb. nov. (Diplobacillus, Allen, Jour. Amer. Vet. Med. Assn., 54, 1918, 307; Diplobacillus, Jones and Little, Jour. Exp. Med., 38, 1923, 139; Hemophilus bovis Hauduroy et al.. Diet. d. Bact. Path., 1937, 247; Moraxella duplex des Bovid^s, Lwoff, Ann. Inst. Past., 62, 1939, 174; Hemophilus ruminaniium Reid and Anigstein, Texas Reports on 592 MANUAL OF DETERMINATIVE BACTERIOLOGY Biol, and Med., 3, 1945, 187.) From Latin hovis, of the ox. Short, plump rods: 0.5 by 1.5 to 2.0 microns, usually occurring in pairs and short chains, with rounded ends. Cap- sulated. Non-motile. Gram-negative. Gelatin: Slow growth at 22°C. Very slow liquefaction. Blood agar colonies : After 24 hours, round, translucent, grayish-white, sur- rounded by a narrow, clear zone of hem- olysis. Deep colonies tiny with a clear hemolytic zone, usually 1.5 mm in diam- eter. After 48 hours, surface colonies somewhat flattened, 3.5 to 4 mm in diameter; deep colonies ellipsoidal and biconvex with hemolytic area of 2.5 to 3 mm in diameter. Blood agar slants: After 24 hours at 38°C, heavy, viscid, grayish-white growth . Coagulated serum liquefied. Broth: Slow growth. Slight turbidity. Considerable sediment. Litmus milk: Alkaline. Partial co- agulation. Indole not produced. Potato: No growth. No acid from glucose or other carbo- hydrates. Not pathogenic for laboratory animals. Killed at 58° to 59°C in five minutes. Aerobic. Source : From cases of acute ophthalmia (pink eye) of cattle. Habitat : In the exudate from cases of acute ophthalmia of cattle. The prob- able cause of bovine infectious keratitis (Baldwin, Amer. Jour. Vet. Res., 6, 1945, 180). Appendix: Other species placed in the genus Moraxella are as follows : Moraxella josephi Lwoff. (Bacillus duplex josephi Scarlett, Annales d'Ocu- listique, 153, 1916, 100 and 485; Lwoff, Ann. Inst. Past., 62, 1939, 171; Moraxella duplex josephi Lwoff, ibid., 174; Bacillus josephi Audureau, Ann. Inst. Past., 64, 1940,126.) Gram-positive. Pathogenic. From the conjunctiva of man. Moraxella Iwoffi Audureau. (Ann. Inst. Past., 64, 1940, 150.) Two varie- ties are recognized : var. bacteroides and var. brevis. From various types of conjunctivitis in man. Moraxella non liquefaciens Lwoff. (Bacterium duplex-nonliquefaciens Oli- ver and Wherry, Jour. Inf. Dis., 28, 1921, 342; Bacillus duplex non-lique- jaciens Hewlett, Med. Res. Council Syst. of Bact., 2, 1929, 418; Lwoff, Ann. Inst. Past., 62, 1939, 171; Moraxella duplex non liquefaciens Lwoff, ibid., 174; Bacilhis duplex non liquefaciens Audureau, Ann. Inst. Past., 64, 1940, 126; Moraxella duplex var. non liquefa- ciens Audureau, ibid., 144.) From an ulcer of the cornea, and from bronchial sputum in man. Genus III. Noguchia Olitsky, Syverton and Tyler.* (Jour. Exp. Med., 60, 1934, 382.) Named for Noguchi, the bacteriologist who iso- lated the type species. Small, slender. Gram-negative rods present in the conjunctiva of man and animals affected by a follicular type of disease ; mucoid type of growth which on first isolation takes place with some difficulty in ordinary media; motile, flagellated, and encap- sulated; aerobic and facultative anaerobic; optimum temperature for growth 28° to 30°C. The type species is Noguchia granulosis (Noguchi) Olitsky, Syverton and Tyler. Key to the species of genus Noguchia. I. Acid from carbohydrates. A. Acid from raffinose, maltose and salicin. 1. Noguchia granulosis. Arranged by Prof. CD. Kelly, McGill University, Montreal, October, 1938. FAMILY PARVOBACTERIACEAE 593 B. No acid from raffinose, maltose and salicin. 2. Noguchia simiae. II. Xo acid from carbohydrates. 3. Noguchia cuniculi. 1. Noguchia granulosis (Xoguchi) Olitsky et al. {Bacterium granulosis Xoguchi, Jour. Exp. Med., J^8, Supp. 2, 1928, 21; Olitsky, Syverton and Tyler, Jour. Exp. Med., 60, 1934, 382.) From Latin, granular. Rods : 0.25 to 0.3 by 0.8 to 1.2 microns, motile by means of a single flagellum, usually polar. Pleomorphic. Gram-neg- ative. Xo growth on plain agar or broth. Blood agar plate : Minute round col- onies, shiny, somewhat raised, almost transparent or slightly grayish in 48 hours. Later the colonies increase in size, are grayish opalescent and some- what sticky. Old colonies have a brown- ish or yellowish tint. Semi-solid Leptospira medium : Gray- ish-white, diffuse growth, forming a delicate zone 1 cm deep. Liquid Leptospira medium: Diffuse, slightl}' cloudy growth, with sticky grayish sediment at the bottom of the tube in old cultures. Acid from glucose, fructose, niannose, sucrose, galactose, maltose, salicin, xylose, mannitol, de.xtrin, arabinose, amj'gdalin and lactose. Small amount of acid from raffinose, inulin, rhamnose and trehalose. Xo acid from dulcitol, sorbitol and inositol . Non-pathogenic for rabbits, guinea pigs, rats and mice. Optimum pH 7.8. Temperature relations: Optimum 15" to 30°C. Grows at 37°C. Aerobe, facultative anaerobe. Distinctive characters : Action on car- bohydrates; agglutination reactions; mo- tility at 15°, none at 37 =C. Source : From trachoma of American Indians at Albuquerque, Xew Mexico. Habitat : Regarded by Xoguchi and others as a cause of trachoma in man. Produces a granular conjunctivitis in monkej's and apes. 2. Noguchia simiae Olitsky et al. (Bacterium simiae Olitsky, Syverton and Tyler, Jour. Exp. Med., 57, 1933, 875; Olitsky et al.. Jour. Exp. Med., 60, 1934, 382.) From Latin simia, ape. Slender rods: 0.2 to 0.3 by 0.8 to 1.2 microns, occurring singly, in pairs, in short chains or parallel arrangement of two or three, having pointed ends. Cap- sules are found. Actively motile by means of a single, rarely a double, flagel- lum, usually polar. Gram-negative. Gelatin plates : Colonies more mucoid and raised than on agar. Gelatin stab : Arachnoid growth along line of inoculation. Xo liquefaction. Agar plates: Small, circular, grayish, translucent, smooth, convex, slightly raised colonies having a sticky or mucoid consistencj'. Blood agar plates : More highly translu- cent and colorless in early growth than on plain agar, becoming grayish after two or three daj's. Agar slants: Grayish-white to white, moist, mucoid, raised, glistening growth. Growth is more profuse when blood is added. Leptospira medium : Homogeneous, dense growth in a 0.5 cm sharpl}^ defined layer, -nith a slight, nebulous, uniform opacity about 1 cm below. In three or four days the lower layer becomes more dense and in time extends to the bottom of the tube. Broth : Uniform turbidit}', with a slight grayish sediment and no pellicle. Litmus milk: Unchanged. Potato : Light tan, spreading, abundant growth. Indole not formed. Xitrites not produced from nitrates. Acid but no gas from glucose, fructose. 594 MANUAL OF DETERMINATIVE BACTERIOLOGY mannose, galactose, xylose, arabinose and rhamnose. Small amount of acid from dextrin. Some strains produce a small amount of acid from sucrose, lac- tose, inulin and mannitol. Raffinose, salicin, dulcitol, amygdalin, maltose, trehalose, sorbitol and inositol un- changed. Serological reactions: Rabbit anti- serum is specific for all strains and no cross agglutination with Noguchia gran- ulosis. Temperature relations : Optimum 28° to 30°C. Thermal death point 56°C for thirty minutes. Aerobe, facultative anaerobe. Distinctive characters : Action on car- bohydrates; agglutination reactions. Source : From inflammatory type (Type II) of spontaneous conjunctival follicu- losis in Macacus rhesus monkeys. Habitat : Causes conjunctival follicu- losis in Macacus rhesus monkeys. 3. Noguchia cuniculi Olitsky, Syver- ton and Tyler. (Jour. Exp. Med., 60, 1934, 382. ) From Latin cuniculus, rabbit . Slender rods: 0.2 to 0.3 by 0.5 to 1.0 micron with pointed ends. Capsules are formed of much finer texture than those surrounding Noguchia granulosis or Noguchia simiae. Actively motile with peritrichous flagella. Non-acid- fast. Pleomorphic forms sometimes noted. Gram-negative. Gelatin agar plates: Grayish, mucoid and confluent colonies. Gelatin stab: Tenuous, arborescent, non-spreading growth. No liquefaction. Agar plates: Small, spherical, translu- cent, slightly grayish, smooth, some- what convex, moist and mucoid colonies with entire edges. Blood agar plates: More profuse, more grayish and less translucent than on plain agar. Agar slants: Slightly grayish, translu- cent, coalescent, glistening, mucoid, homogeneous and non-spreading growth. The water of syneresis appears uniformly cloudy or milkj^ depending on amount of growth. Leptospira medium: After 24 hours, a faint, nebulous surface growth followed by an ingrowing sac -like mass, with its base 5 mm across, lying at the center of the under surface and extending for 5 mm into the medium. The area spreads laterallj^ until at about two or three days there is a uniform, opaque, whitish layer about 1 cm thick which progresses slowly until the bottom of the tube is reached in about seven days. Broth : Uniform turbidity, without pellicle. Litmus milk: Unchanged. Potato : Faint, buff-colored (changing to brown after five days), non-spreading, sparse surface growth. Indole not produced. Nitrites not produced from nitrates. No acid or gas from glucose, fructose, inannose, mannitol, sucrose, raffinose, inulin, galactose, maltose, salicin, xylose, dextrin, arabinose, amygdalin, lactose, dulcitol, rhamnose, trehalose, sorbitol or inositol. Serological relations : Rabbit antiserum is specific for all strains, and no cross agglutination with Noguchia granulosis or Noguchia simiae. Temperature relations : Optimum 28° to 30°C. Thermal death point 56°C for 15 to 30 minutes. Aerobe, facultative anaerobe. Distinctive characters : No action on carbohydrates; peritrichous flagella; ag- glutination. Source : From spontaneous conjunctival folliculosis, Type II of rabbits. Habitat : Causes conjunctival follicu- losis in rabbits. Genus IV . Dialister Bergey el al.* (Manual, 1st ed., 1923, 271.) Minute rod-shaped cells, occurring singly, in pairs and short chains. Non-motile. Rearranged by Prof. D. H. Bergey, Philadelphia, Pennsylvania, 1933. FAMILY PARVOBACTERIACEAE 595 Strict parasites. Growth occurs only under anaerobic conditions in media containing fresh, sterile tissue or ascitic fluid. The type species is Dialister ■pneumosintes (Olitsky and Gates) Bergey et al. 1. Dialister pneumosintes (Olitsky and Gates) Bergey et al. {Bacterium pneumosintes Olitsky and Gates, Jour. Exp. Med., 33, 1921, 713; ibid., 35, 1922, 813; Bergey et al., Manual, 1st ed., 1923, 271; Bacillus pneumosintes Ford, Textb. of Bact., 1927, 634.) From Greek pneu- mon, lung and sintor, murderer or devastator. \evy short rods : 0.15 to 0.3 (in glucose broth 0.5 to 1.0) micron in length, occur- ring singly and occasionally in pairs, short chains or masses. The ends are rather pointed. Xon -motile. Gram- negative. Blood agar colonies : Small, clear, circu- lar, entire, translucent. Growth occurs in media containing fresh sterile rabbit kidney and ascitic fluid. Under strict anaerobic conditions good growth on rabbit blood glucose agar plates. Glucose broth in which Escherichia coli or Bacillus viesentericus (non-spore stage) has grown favors growth. Acid but no gas from glucose. Neither acid nor gas from maltose, lactose, sucrose, inulin or mannitol. Passes Berkefeld Vand N filters. Optimum pH 7.4 to 7.8. No growth at pH 7.0 or pH 8.0. Optimum temperature 37 °C. Does not survive 56 °C for half an hour. Pathogenic for rabbits and guinea pigs. Strict anaerobe. Source : From filteretl nasopharyngeal secretions from influenza patients in the early hours of the disease. Habitat : Nasopharyngeal washings of man. 2. Dialister granuliformans (Pavlovic) Bergey et al. {Bacterium granulifor- mans Pavlovic, Cent. f. Bakt., I Abt., Orig., 112, 1929, 432; Bergey et al., Man- ual, 4th ed., 1934, 341.) From Latin, forming granules. Small rods: Non-motile. Gram-nega- tive. Agar colonies : Very small, transparent. No gas. Broth: Turbid. Litmus milk: Unchanged. Indole not formed. Acid from glucose, sucrose and man- nitol. Passes through Chamberland Lo filters. Pathogenic for rabbits. Optimum temperature 37°C. Anaerobic to microaerophilic. Source : From respiratory tract in influenza. Habitat : Mucous membrane of respira- tory tract. Appendix, Family Parvobacteriaceae:* De Bord (Iowa State Coll. Jour. Sci., 16, 1942, 471) describes a new tribe, Mimeae, which may belong in this fam- ily. The tribe includes three genera : Mima with the species Mima polymorpha and the variety Mima polymorpha var. oxidans; Herellea with the single species Herellea vaginicola; and Colloides with the single species Colloides anoxydana. The organisms are Gram-negative, pleo- morphic, motile or non-motile rods, often showing bipolar staining, and were isolated from the normal vagina and from cases of vaginitis and conjunctivitis. Deacon (Jour. Bact., ^9, 1945, 511) classi- fies nineteen cultures in these genera. * Arranged by Dr. A. Parker Hitchens, University of Pennsjdvania, Philadelphia, Pa., March, 1946. 59G MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY XII. BACTERIACEAE COHN.* (Arch. f. path. Anat. u. Physiol., 55, 1872, 237.) liod-shaped cells without endospores. Motile or non-motile. Gram-positive and Gram-negative. Metabolism complex, amino acids being utilized, and generally carbohydrates. This is a heterogeneous collection of species whose relationships to each other and to other groups are not clear. Only a single genus is recognized at this time. Genus I. Bacterium Ehrenberg. (IV. Evertebrata, Berlin, 1828, 8.) The original description of this genus follows: Bacterium, Novum Genus, Familia Vibrionorum. Character Generis: Corpus polygastricum? anen- terum? nudum, oblongum, fusiforme aut filiforme, rectum, monomorphum (contractione nunquam dilata- tum), iiarum flexile (nee aperte undatiim), transverse in multas partes sponte dividuum. This may be translated as follows : Bacterium, new genus. Family of Vibriona. Character of the genus: Body with many stomachs? without an intestine? naked, oblong, spindle-shaped or filiform, straight, monomorphic (in contraction never dilated), not very pliant (and not definitely wavy), freely separated transversely into many parts. The tj'pe species is Bactermm triloculare Ehrenberg. The original description of this species follows; B. triloculare nov. spec; distincte triloculare s. triarticulatum, subfusiformum, hyalinum. Animalculum 1/300 lineae longum, corpore tereti. Articuli s. septa interna divisionem instantem multi- pliceni transversam indicare videntur. Mobile sed pigrum animalculum. In Oasi Jovis Hammonis Siwae observatuni, praeterea nuUibi. Bacterii Generis jihysiologia huiusque obscurra. Cibo colorato Ventriculos replere hae formae respuunt ideoque ad Polygastrica non misi dubitanter et interim coUocantur. This may he translated as follows; B. triloculare new. spec. Definitely with three compartments or three jointed, subfusiform, hyaline. Animalcules 1/300 of a line in length, with a smooth body. The joints or internal septa are observed to de- velop preliminary to m.ultiplc transverse splitting. A motile but sluggish animalcule. Observed in the Oasis of Jupiter Ammon of Siwa, nowhere else. The physiology of the genus Bacleiium is thus far obscure. These forms refuse to fill their stomachs with colored food and for this reason they are placed with hesitancy and onlj' temporarily in the Polygastrica. The original descriptions are taken from Buchanan, General Systematic Bac- teriology, 1925, 213, and the translations are also furnished by him. Buchanan in his book gives an excellent summary of the nomenclatural status of the term Bacterium on pages 213-230. Since neither the genus nor the type species is characterized in a way to permit definite identification, the term Bacterium is used to cover species of non-spore- forming, rod-shaped bacteria whose position in the system of classification is not definitely established (Breed and Conn, Jour. Bact., 31, 1936, 517). * Completely rearranged by Prof. Robert S. Breed and Mrs. Eleanore JTeist Clise, New York State Experiment Station, Geneva, New York, May, 1946. FA!\ULY BACTERIACEAE 597 Bacterium triloculare Ehrenberg. (Ehrenberg, IV. Evertebrata, Berlin, 1828, 8; Bacillus ehrenhergii Trevisan, I generi e le specie delle Batteriacee, 1899, 18; Bac- terium ehrenbergii De Toni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1022; Bacterium lineola Cohn, Beitrjige z. Biol. d. Pflanzen, /, Heft 2, 1872, 170.) Cohn also regards Vibrio lineola Miller, 1786 and Vibrio lineola {Bacillus lineola, Bactrium lineola) of other authors as synonyms of Bacterium triloculare as explained by Buchanan {Joe. cit., 213 and 521). PVom Latin tri, three and loculus, cells or compartments. Key to the remaining species of genus Bacterium. I. Gram-positive. A. Non -motile. 1. Nitrites produced from nitrates. 1. Bacterium erythrogenes. 2. Bacterium subrufum. 3. Bacterium linens. 4. Bacterium mycoides. 5. Bacterium, mutabile. 6. Bacterium qualis. 2. Nitrites not produced from nitrates. a. Grow on ordinary media. 7. Bacterium racemosum. 8. Bacterium healii. 9. Bacterium insectiphilium. 10. Bacterium tegumenticola . 11. Bacterium minutaferula . 12. Bacterium fulvum. aa. Grow only on sea water media on fresh isolation. 13. Bacterium marinopiscosus. 14. Bacterium sociovivum. 15. Bacterium iinmotum. 3. Action on nitrates unknown. 16. Bacterium ammoniagcn.es . 17. Bacterium minutissimum . B. Motile in young cultures. 1. Nitrites not produced from nitrates. 18. Bacterium incertum. 19. Bacterium imperiale. C. Motile. Proteus-like growth on media. 1. Nitrites not produced from nitrates. 20. Bacterium zopfii. 21. Bacterium zenkeri. Appendixes I and II : These list 34 additional species of Gram-positive, motile or non-motile, non-spore-forming, rod-shaped bacteria. See p. 609 and 612. II. Gram-negative. Digest cellulose. Do not digest agar. A. Non-motile. Gelatin liquefied. Chromogenic. 1. Milk acid. 22. Bacterium idoneum. 22a. Bacterium liquatum. 598 MANUAL OF DFTERMIXATIVE BACTKRIOLOGY B. Non -motile. Gelatin liquefied. Non-chromogenic. 1. Milk acid. a. Ammonia produced; indole not formed. 23. Bacterium udum. C. Non-motile. Gelatin not liquefied. Non-chromogenic. 1. Milk unchanged. a. Ammonia not produced; indole not formed. 24. Bacterium lucrosum. 2. Milk acid. a. Ammonia not produced; indole not formed. 25. Bacterium acidulum. 26. Bacterium castigatum. D. Motile. Gelatin liquefied. Chromogenic. 1. Milk acid. a. Ammonia produced; indole is formed. 27. Bacterium hibulum. Appendices I to III: These list additional species of cellulose-digesting, Gram -negative, usualh' motile, rod-shaped bacteria. See p. 615 and 622. Also similar species that utilize bacterial poh'saccharides as a sole source of carbon. Seep. 623. III. Gram-negative. Digest agar. A. Non-motile. 1. Nitrites not produced from nitrates. a. Acid from glucose and other sugars. 28. Bacteriuttt nrnckii. aa. Do not form acid from glucose. 29. Bacterium polijsiphojiiae. 30. Bacterium drohachense. 2. Action on nitrates unknown. a. Do not form acid from glucose. 31. Bacterium delesseriae. 32. Bacterium boreale. 33. Bacterium ceramicola. B. Motile but position of flagella not given. May be either peritrichous or polar. 1. Nitrites not produced from nitrates. 34. Bacterium rhodomelae. 2. Action on nitrates unknown. 35. Bacterium alginovorum. 36. Bacterium fucicola. Appendices I and II: These describe 9 additional species and list others that digest agar. All are Gram-negative, motile or non-motile rod-shaped bacte- ria. See p. 627. IV. Gram-negative. Digest chitin. A. Motile but position of flagella not given. 1. Non-chromogenic. 37. Bacteriuin chititwphilum. 2. Yellow chromogenesis. 38. Bacterium chitinochroma. FAMILY BACTERIACEAE 599 Appendix I: One additional species is described. See p. 632. V. Gram-negative. Phosphorescent bacteria. A. Non -motile coccobacilli from sea water. 1. No liquefaction of gelatin. 39. Bacterium phosphoreum. B. Motile rods from sea water. Position of flagella not given. 1. Xo growth in broth, and on coagulated blood serum or potato. 40. Bacterium phosphorescens indigenus. C. Xot stated whether motile or non-motile. From diseased insect larvae. 1. Yellow growth on potato. 41. Bacterium hemophosphoreum. Appendix I : This includes a list of more than 40 additional so-called species of phosphorescent l)acteria. See p. 634. VI. Gram-negative. Facultative autotrophic bacteria which secure energy from the oxidation of hydrogen and utilize carbon from CO-. A. Xon-motile. 1. Growth shows a red chromogenesis. 42. Bacterium erythrogloeum. B. Motile with peritrichous flagella. 1. Yellow chromogenesis. 43. Bacterium lentulum. 2. Ivory-colored colonies. 44. Bacterium leucogloeum. VII. Gram-negative. Plant pathogens. A. X' on-motile. 1. Gelatin not licjuefied. 45. Bacterium stewartii. B. Motile with a polar flagellum. 1. Gelatin not liquefied. a. Colonies mustard yellow on agar. 46. Bacterium tardicrescens. b. Colonies hone}' to Xaples yellow on agar. 47. Bacterium albilineans. Appendix I : This includes 19 additional species placed in Bacterium or Bacil- lus by their authors. All are reported to cause or to be associated with plant disease. See p. 639. VIII. Gram-negative. Miscellaneous species. A. Produce a pink to red chromogenesis. 1. Motile. a. Gelatin not liquefied. 48. Bacterium rubefaciens. aa. Gelatin liquefied. 49. Bacterium rubidum. 2. Xon-motile. a. Gelatin not liquefied. 50. Bacterium later iccum. 600 MANUAL OF DETERMINATIVE BACTERIOLOGY B. Do not produce pink or red chromogenesis. 1. Motile. a. Produce clouding in alginic acid liquid medium, b. From sea water. 51. Baclerinm alf/inicum. bb. From soil. 52. Bacterium terrestrnhiinicum. aa. Action on alginic acid unknown. b. Causes a disease of swans. 53. Bacterium cijqni. 2. Non -motile. a. Causes red spot disease of carp. 54. Bacterium ci/prinicida. aa. Causes liberation of ammonia from a mixture of horse manure and urine. 55. Bacterium, parvulum. aaa. Utilizes formates in a litiuid medium with the formation of a red- dish pellicle. 56. Bacterium methylicum. Appendix I: Miscellaneous described species of non-spore-forming bacteria placed by their authors in the genus Bacillus. See p. 64.3. Appendix II: Includes anaerobic bacteria that produce methane. See p. 645. Appendix III: Miscellaneous species of non-spore-forming bacteria listed but not described. See p. 647. 1 . Bacterium erythrogenes Lehmann and Neumann. (Bacterium lactis ery- throgenes Grotenfelt, Fortschr. d. Med., 7, 1889, 41; Bacillus lactis erythrogenes Sternberg, Manual of Bact., 1893, 636; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 253; Bacillus erythrogenes Matzu.schita, Bakt. Diagnostik, 1902, 220; Corynebacterium erythrogenes Kiss- kalt and Berend, Cent. f. Bakt., I Abt., Orig., 81, 1918, 446; Erythrobacillus erythrogenes Holland, Jour. Bact., 5, 1920, 218; Erythrobacillus (lactis) ery- throgenes Holland, ibid.; Serratia lactica Bergey et al., :\Ianual, 1st ed., 1923, 93; Chromobacterium lactis erythrogenes Top- ley and Wilson, Princip. Bact. and Im- mun., 1, 1931, 402.) From Greek, red- producing. Micrococcus Lactis erythrogenes Conn, Esten and Stocking, Ann. Rept. Storrs (Conn.) Agr. E.xp. Sta., 18, 1906, 117 is stated to be allied to if not identical witli the above species. Reds: 0.3 to 0.5 by 1.0 to 1.4 microns, in broth often up to 4.3 microns long, occurring singly, and having rounded ends. Non-motile. Stain with the usual aniline dyes. Gram-positive (Leh- mann and Neumann, loc. cit.). Gelatin colonies: Small, circular, gray- ish, becoming yellow, sinking into the medium. Crateriform liquefaction. Yellow sediment. Medium becomes rose-colored. Gelatin stab : Surface growth a whitish, later yellow, circular, thin layer. Weak growth in stab. Slow liquefaction at the surface, the liquid becoming red, with yellow sediment. The solid portion as- sumes a weak rose color. Agar stab: Moist, fairly luxuriant, yellow growth, the medium assuming a rose to wine color. Broth: Turbid, yellow. Pellicle (Ful- ler and Johnson, Jour. Exp. Med., 4, 1899, 609). Litmus milk: Acid. Slow coagulation, htiving a clear fluid which becomes blood- red in color. Reaction becomes alkaline. Sterile milk: Casein slowly precipi- tated, later peptonized. Reaction neu- FAMILY BACTERL\CEAE 601 tral or alkaline A stratum of blood-red serum is seen above the precipitated casein and above this a yellowish-white layer of cream. An intensive sweet odor that becomes disagreeable. Potato: Growth rapid, spreading, grayish, later yellow. On incubation a deep golden yellow color develops after 6 to 8 days. A darkening of the medium occurs around the culture, but soon dis- appears ; later the whole potato becomes a weak yellowish -red. Indole not formed (Fuller and John- son, loc. cit.). Indole formed (Chester, Manual Determ. Bact., 1901, 174). Blood serum: Liquefied (Fuller and Johnson, loc. cit.). Not liquefied (Heff- eran, Cent. f. Bakt., II Abt., 11, 1903, 456). Nitrites produced from nitrates. No gas from carbohydrates. Slight H;S production (Matzuschita, loc. cit.). Red pigment insoluble in water, alco- hol, ether, chloroform, and benzol. Soluble (Hefferan, loc. cit., 529) . Yellow pigment insoluble. Distinctive character: Milk becomes blood-red in 12 to 20 days. Non-pathogenic for mice (Fuller and Johnson, loc. cit.). Optimum temperature 28'' to 35°C. Aerobic (Fuller and Johnson, loc. cit.). Facultative anaerobe CHefferan, loc. czi., 530). Source: Isolated from red milk bj- Hueppe in Wiesbaden in 1886. Isolated from feces of a child by Baginsky (Cent, f. Bakt., 6, 1889, 137). Isolated from Ohio River water by Fuller and Johnson (loc. cit.). Isolated from Mississippi River water bj' Hefferan (Joe. cit.). Tataroff isolated a rose fluorescent coc- cobacterium {Bacillus rosafluorescens Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., ^, 1896, 305; Bacterium rosa jluorescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 142) which Migula reports as identical, but which Hefferan considers atypical. Habitat : Probably widely distributed in nature. 2. Bacterium subrufum Burri and Staub. (Burri and Staub, Landwirtsch. Jahrb. d. Schweiz, Jfi, 1926, 1006; Ser- ratia subrufa Bergey et al.. Manual, 3rd ed., 1930, 123.) From Latin sub, some- what and riifus, red. This organism is stated to be closelj^ related to or possibly identical with Bacterium eryihrogenes. 3. Bacterium linens Weigmann. (Or- ganismus IX, Wolff", Milchwirt. Zent., 5, 1909, 145; Weigmann, in Wolff, Cent. f. Bakt., II Abt., 28, 1910, 422, and in Weigmann, ^Mykologie der Milch, 62, 1911,220.) From Latin, daubing, smear- ing, or spreading over. Also see Steinfatt, Milchwirt. Forsch., 9, 1930, 7; Kelly, Jour. Dairy Sci., 20, 1937, 239; Albert, Long and Hammer, Iowa Agr. Exp. Sta. Res. Bui. 328, 1944. Rods: Average 0.62 bj' 2.5 microns when grown 1 to 2 days on trj^ptone glu- cose extract agar. Non -motile (Wolff). Gram-positive (Kelly, loc. cit.). Gelatin colonies: At 18°C punctiform at first; after 12 days about 1 mm in diameter, compact, circular, shiny, brownish-j-ellow to red-brown. Lique- faction. Gelatin stab: At 21°C crateriform liquefaction, becoming infundibuliform on extended incubation. Rate of lique- faction varies considerably with differ- ent cultures, some completing it in 15 daj's, others not completing it even on long incubation. Agar colonies: On tryptone glucose extract agar at 21 °C after 1 to 2 days, colonies convex, glistening, entire and cream-colored, becoming brown on ex- tended incubation; diameters 2 to 5 mm. On special cheese agar with incubation in oxygen, luxuriant growth, the color becoming bright orange to reddish- brown in 4 or 5 days. 602 MANUAL OF DETERMINATIVE BACTERIOLOGY Agar stab: Heavy surface growth on tryptone glucose extract agar at 21°C with no growth along the line of inocu- lation. Agar slant: On tryptone glucose ex- tract agar at 21 °C after 2 days growth abundant, glistening, filiform, non -viscid and cream-colored. After extended in- cubation the color usually is brown. On special cheese agar in an atmosphere of oxygen the growth is bright orange to reddish-brown in 4 or 5 days. Broth: Turbidity and sediment. Potato : At 21°C after 5 days, growth is scanty, smooth, glistening, and varies in color from grayish to brownish -orange. Litmus milk: At 21°C the changes are very slow. After 6 or 7 days the reac- tion becomes alkaline and a yellow sedi- ment appears. After approximately 10 days some digestion is evident, complete digestion generally requiring several weeks to over a month. A distinct am- moniacal odor, more or less objection- able, produced in old cultures. No coagulation. Ropiness often produced on extended incubation. Indole not produced. Nitrites produced from nitrates. Methyl red and Voges Proskauer reac- tions negative. Hydrogen sulfide produced in broth and on agar by some cultures but not by others. Natural fats not hydrolyzed. No acid or gas from arabinose, dex- trin, glucose, dulcitol, galactose, inulin, lactose, fructose, maltose, mannitol, raffinose, rhamnose, salicin, sorbitol, sucrose or xylose. Ethyl, propyl, but3i and amyl alco- hols o.xidized largely to corresponding acids ; he.xyl and heptyl alcohols attacked much less actively. Catalase rapidly produced in or on various media. Aerobic. Growth temperatures: Growth at 8° and 37°C but not at 45°C, with the opti- mum at about 21 °C. Heat resistance low, cultures being killed at 62.8°C in a few minutes. Growth in the pH range 6.0 to 9.8; no growth at pH 5.0 or below. Salt tolerant, cultures growing readily in a concentration of 15 per cent salt in broth or skim milk, with certain cultures apparently capable of growing somewhat in much higher concentrations. Closely related to or identical with Bacterium erythrogenes Lehmann and Neumann. Source: Originally isolated by Wolff from the surface flora of various soft cheeses. Habitat: Widely distributed in and especially on the surface of dairy prod- ucts including blue, brick, camembert, limburger, oka and cheddar cheeses, butter, milk and cream. Also found in various feeds including grains, silage, green plants, hay and straw, and in water, soil, manure, and air. 4. Bacteriixm mycoides (Grotenfelt) Migula. (Bacterium mycoides roseum Grotenfelt, Fortschr. d. Med., 7, 1889, 46; Bacillus mycoides roseus Sternberg, Manual of Bact., 1893, 640; Migula, Syst. d. Bakt., 2, 1900, 482; Bacillus mycoides-roseus Holland, Jour. Bact., 5, 1920, 219; Erythrohacillus mycoides- roseus Holland, ibid.; Serratia rosea Bergey et al.. Manual, 1st ed., 1923, 96; Chromohacteriuvi mycoides roseum Top- ley and Wilson, Princip. of Bact. and Immun., ?, 1931, 402.) From Greek mykes, fungus and eidos, form. Rods: Non -motile. Gram-positive. (ielatin colonies: Red, felt-like. Liquefaction. Gelatin stab: Rapid liquefaction. Red pellicle. Red sediment. Colonies composed of interlacing fila- ments (Crookshank, Textb. of Bact. and Inf. Dis., 1900, 524). .\gar stab: Red color produced if grown in dark; a white color in presence of light. FAMILY BACTERIACEAE 603 Optimum temperature: Room tem- perature. Pigment soluble in water. Distinctive characters: Morphologi- cally like the anthrax bacillus. Appear- ance in gelatin. Production of a brilliant rose color when grown in the dark; colonies grown in the light are white, but they assume the red color if developed further in the dark. Source: Isolated from Wiesbaden soil by Scholl. Habitat: Unknown. Note: It has been claimed that this or a similar organism forms spores (Matzuschita, Bact. Diag., 1902, 168; Perlberger, Cent. f. Bakt., II Abt., 62, 1924, 8). However cultures of Scholl's organism received from the Krai collec- tion by Hefferan (Cent, f . Bakt., II Abt., 11, 1903, 458) and by Breed in 1926 (per- sonal communication) did not form spores. These cultures produced ni- trites from nitrates and failed to liquefy gelatin. 5. Bacterium mutabile Steinhaus. (Jour. Bact., 42, 1941, 775.) From Latin mutabilis, changeable. Short rods: On agar, 0.7 to 0.9 by 1.0 to 2.0 microns. In fluid media, such as tryptophane broth, pleomorphic, bi- zarre forms frequently appearing slightly branched. Non -motile. Gram-posi- tive. Gelatin stab: Very slow liquefaction. Agar colonies: Cream to yellow, cir- cular, smooth, glistening, opaque. Broth : Moderate turbidity, slight sedi- ment. Litmus milk: Alkaline, soft curd, slow peptonization. Indole not produced. Hydrogen sulfide not produced. Nitrites produced from nitrates. Starch not hydrolyzed. Glucose, lactose, sucrose and maltose not fermented. Aerobic. Source: From the alimentary tract of the lyreman cicada, Tibicen linnei Smith and Grossbeck. Habitat: Unknown. 6. Bacterium qualis Steinhaus. (Jour. Bact., 42, 1941, 774.) From Latin qualis, of what kind. Short rods: Very short on solid media, frequently ellipsoidal in shape. In fluid media: 0.5 to 0.7 by 1.4 to 2.2 microns, occurring singly. Non-motile. Gram- positive. Gelatin stab: Liquefaction. Agar colonies: Small (1 mm), white, glistening, transparent, circular, entire. Agar slant: Filiform, smooth, glis- tening. Broth: Almost clear; slight turbidity in serum and glucose broth. Litmus milk: No change. Indole not produced. Hydrogen sulfide not produced. Slight production of nitrites from ni- trates. Starch not hydrolyzed. Acid from glucose, sucrose and mal- tose. Lactose not fermented. Source: From the alimentary tract of the tarnished plant bug, Lygus praten- sis L. Habitat: Unknown. 7. Bacterium racemosum Zettnow. (Zettnow, Cent. f. Bakt., I Abt., Orig., 77, 1915, 209; Zetlnowia racemosa Ender- lein, Bakt. Cyclogenie, Berlin, 1925,259; Flavobacterium racemosum Bergey et al.. Manual, 1st ed., 1923, 115.) From Latin race7iwsus, branching. Filaments: 0.5 to 0.8 by 10 to 12 mi- crons. Branching forms found. Non- motile. Gram-positive. Gelatin colonies: White, circular, soft, granular, brownish, entire. Gelatin stab: White surface growth. Liquefaction napiform. Agar slant: Light yellow, limited growth. Broth: Turbid. 604 MANUAL OF DETERMINATIVE BACTERIOLOGY Litmus milk: Coagulated, becoming alkaline. Potato: Dirty -yellowish, limited streak. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 20°C. This species is selected as the type species for the genus Zcitnowia Ender- lein {loc. cit.). Source: Contamination on agar plate. Habitat: Unknown. 8. Bacterium healii Buchanan and Hammer. (Buchanan and Hammer, Iowa Agr. Exp. St a. Research Bull. 22, 1915, 249; Escherichia healii Bergey et al., Manual, 1st ed., 1923, 200; Achru- mohacter healii Bergey et al., Manual, 2nd ed., 1925, 157.) Rods: 0.5 to 0.7 by 2.2 to 12.9 microns, occurring singly and in short chains. Non-motile. Gram-positive. Gelatin stab: Stratiform litiuefaction. Villous growth in stab. Agar colonies: Large, white, rhizoid. Agar slant: White, hard. growth, with no tendencj' to stringiness. Broth: Gray pellicle and sediment. Litmus milk: Slightly acid, becoming slimy, coagulated, peptonized. Potato: Heavy, white, glistening growth. Indole not formed. Nitrites not produced from nitrates. Acid without gas from glucose, fruc- tose, maltose, sucro.se, salicin and starch. No acid from mannitol, lactose, raffinose or inulin. Aerobic, facultative. Optimum temperature 22°C. Source: Slimy milk. Habitat : Unknown. 9. Bacterium insectiphilium Steinhaus. (Jour. Bavi.. 42, 1941, 777.) From M. L. insect, insect and Greek philos, loving. Rods: 0.8 to 1.2 by 1.0 to 2.8 microns, occurring singly. At times appearing almost as cocci or coccobacilli. Non- motile. Gram -positive. Gelatin stab: Liquefaction. Agar colonies: Light greenish-yellow, circular, entire, raised, glistening, smooth, opaque. Agar slant: Filiform, raised, smooth, glistening, opaque growth. Broth: Moderate turbidity, slight vis- cid sediment. Litmus milk: Alkaline, peptonization, and slow reduction. Potato: Greenish-yellow, thick, moist growth . Indole not produced. Nitrites not produced from nitrates. Hydrogen sulfide not produced. Starch slightly hj-drolj'zed. No action on the following carbohy- drates: Glucose, lactose, sucrose, mal- tose, fructose, mannitol, galactose, arab- inose, xylose, dextrin, salicin, rhamnose, raffinose, trehalose, sorbitol, inulin, dul- citol, glycerol, adonitol, mannose. Aerobic. Source: From the body wall of the bagworm, Thyridopteryx ephemeraefor- juis Haw. Habitat : Unknown. 10. Bacterium tegumenticola Stein- haus. (Jour. Bact., ^2, 1941, 775.) From Latin legumenlum, cover, skin and cola, dweller. Small rods: 0.5 to 0.8 by 1.0 to 1.5 mi- crons. Have a tendency to be ellipsoidal on solid media. Non-motile. Gram- positive. Gelatin stab: Generally no liciuefac- tion. ^'ariable. Agar colonies: Tiny (1 nmi), white, convex, glistening, circular, entire. Agar slant: Filiform, glistening, gray- ish-white growth. Broth: Slight turbidity; sediment. Litmus milk: No change. Indole not produced. Hydrogen sulfide not produced. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid slowly produced from glucose FAMILY BACTERIACEAE 605 and maltose. Acid from sucrose. Lac- tose not fermented. Source: From the integument of the bed-bug, Cimex lectularius L. Habitat: Unknown. 11. Bacterium minutaferula Steinhaus. (Jour. Ba.ct.,43, 1941, 778.) From Latin, small rod. Very small rods: 0.4 to 0.9 by 0.7 to 1.0 micron, occurring singly. Xon-motile. Gram-positive. Gelatin stab: Xo liquefaction. Agar colonies: Colorless to faint gray, circular, smooth, entire, glistening. Agar slant: Very thin, transparent, glistening growth. Broth: Slight turbidity and sediment. Litmus milk: Xo change at first; slightly acid after one week. Indole not produced. Hydrogen sulfide not produced. Xitrites not produced from nitrates. Starch not hydrolyzed. Acid from glucose after 4 days . Slight acid from sucrose. Lactose and maltose not fermented. Aerobic. Source: From triturated specimen of the mud-dauber wasp, Sceliphron ce- mentarium Dru. Habitat: Unknown. 12. Bacterium fulviun (Zimmermannj Chester. {Bacillus fulvus Zimmermann, Bakt. unserer Trink- u. Xutzwasser, Chemnitz, 1, 1890, 44; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., .9, 1897, 107; Flavohacterium julvum Bergey et al., Manual, 1st ed., 1923, 115.) From Latin fulvus, dull yellow. Rods: 0.8 bj^ 0.9 to 1.3 microns, occur- ring singly, in pairs and in chains. Xon- motile. Gram-positive. Gelatin colonies: Circular, convex, reddish -yellow. Gelatin stab: Convex, reddish-yellow surface growth. Good growth in stab. Slow liquefaction. Agar slant: Orange-red, glistening streak. Broth: Turbid with yellow sediment. Litmus lactose broth: Acid, or acid then alkaline (Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 368). Potato: Slowly spreading, j-ellowish, glistening growth. Indole formed (Dj'ar, loc. cit.). Xitrites not produced from nitrates (Bergey) . Aerobic, facultative. Optimum temperature 30°C. Source: From Chemnitz and Dobeln tap water (Zimmermann). From dust and water (Dyar) . Habitat: Water. 13. Bacterium marinopiscosus ZoBell and Up ham. (Bull. Scripps Inst. Ocean- ography, La JoUa, 5, 1944, 258.) From Latin marinus, pertaining to the sea, and piscosus, fish. Rods: 1.2 to 1.6 by 2.0 to 4.7 microns, with rounded ends, show granular stain- ing, occurring singly, in pairs and long chains. Xon-motile. Gram-positive, but many cells tend to decolorize leaving Gram-positive granules. All differential media except the fresh- water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: Gray, circular, con- vex, 1 mm. Xo pigment. Gelatin stab: Liquefaction napiform, becoming crateriform to stratiform with age. Complete in 50 days. Agar colonies : 2 to 4 mm, circular, con- vex, entire, smooth, irregular edge. Agar slant : Luxuriant, beaded, glisten- ing, butyrous growth with no pigment. Sea-water broth: Xo turbidity, abun- dant fiocculent sediment, slight surface ring. Fresh-water broth: Good growth. Litmus milk: Decolorized, neutral, top peptonized. Potato: Heavy, white, raised, mucoid, dull growth. Potato darkened. Indole not formed. Xitrites not produced from nitrates. Acid but no gas from glucose and man- 606 MANUAL OF DETERMINATIVE BACTERIOLOGY nitol. No acid from maltose, lactose, sucrose, glycerol, xylose or salicin. Starch is hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Casein is digested. Fats are not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source: Found on the skin of marine fish. Habitat: Not known from other sources. 14. Bacterium sociovivum ZoBell and Upham. (Bull. Scripps Inst. Ocean- ography, La Jolla, 5, 1944, 269.) From Latin socius, associate and vivum, to live. Rods: 0.5 to 0.8 by 3.0 to 4.0 microns, with rounded ends, occurring singly, in pairs, and chains. Non-motile. Gram- positive but tends to destain, leaving Gram-positive cell wall and granules. All differential media except the fresh- water broth, litmus milk and potato were prepared with sea water. Gelatin colonies: Irregular, sunken, filamentous margin, grayish-white. Gelatin stab: Crateriform liquefaction becoming stratiform. Agar colonies: 2 to 4 mm, circular, convex, smooth, entire, darker center. Agar slant: Luxuriant, beaded, glis- tening, butj'rous growth with no pig- ment. Sea-water broth: No pellicle, no tur- bidity, heavy flocculent sediment. Fresh -water broth: Fair growth. Litmus milk: Decolorized, neutral, completely peptonized in 20 days. Potato: Abundant, dull, light cream- colored growth. Potato darkened. Indole not formed. Nitrites not produced from nitrates. Acid but no gas from glucose, maltose, and mannitol. No acid from glycerol, lactose, sucrose, or salicin. Starch is hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Casein is digested. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source: Found associated with seden- tary organisms in the sea. Habitat: Commonly found on sub- merged surfaces and on sessile diatoms in sea water. 15. Bacteritim immotum ZoBell and' Upham. (Bull. Scripps Inst. Ocean- ography, 5, 1944, 271.) From Latin, meaning immobile or stationary. Rods: 0.8 by 3.1 to 8.6 microns, with rounded ends, occurring singly, in pairs, and long chains. Non -motile. Gram- positive but tend to destain leaving Gram-positive outline and granules. All differential media except the fresh- water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies:^ Small, circular, raised, gray, slowly digest gelatin. Gelatin stab: Crateriform liquefaction becoming infundibuliform. Beaded growth along line of stab. No pigment. Agar colonies : 1 to 2 mm, circular, con- vex, smooth, lobate margin, darker centers. Agar slant: Luxuriant, glistening, echinulate, mucoid growth with no pigment. Sea-water broth : No pellicle, moderate turbidit}', abundant, flocculent sedi- ment. Fresh-water broth: Scanty growth. Litmus milk: Decolorized, neutral, partly peptonized in 20 days. Potato: Luxuriant, mucoid, creamy growth which darkens potato. Indole not formed. Nitrites not produced from nitrates. Acid but no gas from glucose, maltose, xylose, and mannitol. No acid from glycerol, lactose, sucrose, or salicin. Starch is hydrol3^zed. Hvdrogen sulfide not formed. FAMILY BACTERIACEAE 607 Ammonia produced from peptone but not from urea. Casein is digested. Fats not hydrolyzed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source: Found associated with marine sedentary organisms. Habitat: Xot known from other sources. 16. Bacterium ammoniagenes Cooke and Keith. (Cooke and Keith, Jour. Bact., 13, 1927, 315; Alcaligenes ammoni- agenes Bergey et al., Manual, 3rd ed., 1930, 367.) From M. L. ammonia and Latin genera, develop. Rods with rounded ends, 0.8 by 1.4 to 1.7 microns, occurring singly. Xon- motile. Gram-positive. Gelatin stab: Xo liquefaction. Agar colonies: Circular, flat, smooth, entire, gray. Agar slant : Growth moderate, smooth, flat, opaque, glistening, butyrous, amor- phous. Broth: Moderate turbidity, with floc- culent sediment. Litmus milk: Slightly alkaline. Indole not formed. Xo action on carbohydrates. Blood serum not liquefied. Urea is fermented forming ammcmia. Aerobic, facultative. Optimum temperature 30°C. Source: From feces of infants. Habitat: Presumably widely dis- tributed in putrefying materials. 17. Bacteriimi minutissimum Migula. (Bacillus pyogenes miriutissimus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 447; Bacterium pyogenes minu- tissimus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 89; Migula, Syst. d. Bakt., 2, 1900, 418; Eberlhella minutissima Bergey et al.. Manual, 1st ed., 1923, 228; Shigella minutissima Bergey et al., Alanual, 3rd ed., 1930, 359.) From Latin, smallest. Description from Kruse {loc. cit.). Description not different from that of Bacillus tenuis sputigenes Pansini, ac- cording to Chester (Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 89). Rods: 0.5 by 1.0 micron, occurring singly and in pairs. Xon-motile. Gram-positive. Gelatin stab: Xo liquefaction. Yel- lowish growth spreading slightly on surface. Growth on agar and blood serum is not characteristic. Acid but no gas from glucose and lac- tose. Xo characteristic odor. Xot pathogenic for mice and rabbits. Aerobic, facultative. Optimum temperature 37 °C. Source: Isolated from a facial abscess. Habitat: Xot known from other sources. 18. Bacteriiun incertum Steinhaus. (Jour. Bact., 42, 1941, 776.) From Latin incertus, uncertain. Short rods: 0.5 to 0.8 by 1.0 to 1.5 microns, occurring singly and occasion- ally in pairs. Young cultures motile, after 48 hours generally non-motile. Gram-positive; after 48 hours many cells become Gram-negative. Gelatin stab: Xo liquefaction. Agar colonies: Tiny, grayish-white, smooth, almost transparent. Does not grow well on nutrient agar. Xorth's gelatin chocolate agar slant: Filiform, thin, transparent growth. Brown color of chocolate medium changes to yellowish-green. Blood agar: Alpha hemolysis at first; after three days beta hemolysis. Broth: Almost clear; ver\- slight growth. Litmus milk: Xo change. Indole not produced. Hydrogen sulfide not produced. Xitrites not produced from nitrates. Starch not hj'drolyzed. Acid but no gas from glucose, sucrose. 608 MANUAL OF DETERMINATIVE BACTERIOLOGY fructose, inannose, and maltose. No fermentation of lactose, rhamnose, galac- tose, mannitol, dulcitol, inositol, or sorbitol. Voges-Proskauer test: Negative. Microaerophilic. Source: From the ovaries of the lyre- man cicada, Tibicen linnei Smith and Grossbeck. Habitat: Unknown. 19. Bacterium imperiale Steinhaus. (Jour. Bact., ^2, 1941, 777.) From Latin imperialis, referring to the imperial moth. Small rods: 0.5 to 0.8 by 1.0 to 1.7 microns, occurring singly and in pairs. A few cells motile in young cultures. Gram-positive. Gelatin stab: No liquefaction. Agar colonies: Circular, entire, almost translucent, pinkish-orange to yellow pigment. Agar slant: Filiform, glistening, opaque growth. Broth: Slight to moderate turbidity; slight sediment. Litmus milk: No change at first, later slightly acid. Potato: Heavy, glistening, moist growth; reddish to yellowish-orange. Indole not produced. Hydrogen sulfide not produced. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid but no gas from glucose, sucrose, maltose, fructose, mannitol, galactose, arabinose, xylose, salicin, raffinose, tre- halose, sorbitol, mannose, adonitol, esculin, and slight acid from lactose and dextrin. Inuliii, dulcitol, glycerol, rhamnose, adonitol, and inositol not fermented. Aerobic. Source: From the alimentary tract of the imperial moth, Eacles ■imperialis Dru. Habitat: Unknown. 20. Bacterixmi zopfii Kurth. (Kurth, Bericht. d. deutsch. Botan. Gesellschaft, /, 1883, 97; Kurthia zopfii Trevisan, Atti della Accad. Fisio-Medico-Statistica in Milano, Ser. 4, 8, 1885, 92; Helikobac- terium zopfii Escherich, Miinch. med. Wchnschr., 33, 1886, 2, quoted from En- lows, U. S. Hygienic Lab. Bull. 121, 1920, 47; Bacterium (Proteus) zopfii Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103; Bacillus zopfii Migula, Syst. d. Bakt., 2, 1900, 815; Zopfius zopfii Wenner and Rettger, Jour. Bact., 4, 1919, 334.) Named for W. Zopf, German botanist. This is the type species of the genus Kurthia Trevisan. (Trevisan, too. cit.; Zopfius Wenner and Rettger, Jour. Bact., 4, 1919, 334.) Rods : 0.8 by 3.5 microns, with rounded ends, occurring in long curved chains. Motile with peritrichous flagella. Gram-positive. Gelatin colonies: Radiate, filamentous, gray. Gelatin stab: Arborescent growth in stab. No liquefaction. Agar colonies: Fimbriate. Agar slant: Spreading, gray, fimbriate growth . Broth: Slow, moderate growth. Litmus milk: No change. Potato: Moderate, gray growth; me- dium becoming dark. No HoS produced. Indole not formed. Nitrites not produced from nitrates. Aerobic, facultative. Optimum temperature 25° to 30°C. Habitat: Decomposing materials. 21. Bacterium zenkeri (Hauser) Ches- ter. (Proteus zenkeri Hauser, Ueber Faulnissbakterien, 1885; Bacillus zenkeri Trevisan, I generi e le specie delle Bat- teriacee, 1889, 17; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103; Zopfius zenkeri Wenner and Rettger, Jour. Bact., 4, 1919, 334; Bacillus pro- teus-zenkeri Holland, Jour. Bact., 5, 1920, 220; Kurthia zenkeri Bergey et al., Manual, 2nd ed., 1925, 215.) Named for K. Zenker, German pathologist. FAMILY BACTERIACEAE 609 Rods: 0.65 by 1.6 to 2.3 microns, occur- ring in pairs and in chains. Motile with peritrichous flagella. Gram-posi- tive. Gelatin colonies: Feather}^, with fila- ments extending in all directions. Gelatin stab: Surface growth like colonies. Xo arborescent growth in stab. Xo liquefaction. Agar colonies: Thin, filamentous, spreading, grayish. Agar slant: Thin, bluish-gray, fila- mentous growth. Broth: Slightly turbid, with gray sediment. Litmus milk: Xo change. Potato: Barely visible, yellowish- gray, glistening growth. Indole not formed. Nitrites not produced from nitrates. Xo H2S formed. Aerobic, facultative. Optimum temperature 30°C. Habitat: Decomposing materials. Note: Wenner and Rettger, loc. cit., consider the last two species to be identical. Appendix I: The following Gram-posi- tive, motile species may belong with the above group. All have been placed at one time or another in the genus Achro- mobactcr or in the genus Flavohacterium. 1. Achromobacter lipolyticum (Hussi Bergey et al. {Bactridium lipolyticum Huss, Cent. f. Bakt., II Abt., 20, 1908, 474; Bergey et al.. Manual, 2nd ed., 1925, 158.) From Greek, fat -dissolving. Small, oval rods: 0.3 to 0.5 by 0.7 to 1.4 microns. Motile, possessing peri- trichous flagella. Gram -positive. Gelatin colonies: Circular, grayish to transparent with irregular margin. Gelatin stab: Infundibuliform lique- faction. Agar colonies: Growth circular, gray, smooth, glistening, with entire margin. Broth : Turbid with granular sediment. Litmus milk: Coagulated, peptonized, becoming alkaline. Potato: Moist, glistening, grajash growth. Indole is formed. Acid from glucose, sucrose, raflSnose, xylose, mannitol and glycerol. Fats are split in milk, giving rise to a rancid odor and a bitter taste. Aerobic, facultative. Optimum temperature 35°C. Source : From the udder of a cow giving abnormal milk. Habitat: Milk. 2. Achromobacter stearophilum (Wein- zirl) Bergey et al. (Bacillus stearo- philus Weinzirl, Jour. Med. Res., S9, 1919, 404; Bergey et al., Manual, 1st ed., 1923, 145.) Rods: 0.8 by 5.0 microns, occurring singly. Motile. Gram-positive. Gelatin colonies: Scanty development. Pumpkin gelatin stab : Filiform growth in stab. X'o liquefaction. Pumpkin agar colonies : Small, smooth, convex, gra}', entire. Pumpkin juice: Slightly turbid. Pumpkin milk: Acid, coagulated. Potato: Slight, smooth, gray, glisten- ing, fUiform growth. Indole not formed. Nitrites not produced from nitrates. No acid from carbohydrate media. Starch from pumpkin hydrolyzed. Aerobic, facultative. Optimum temperature 20°C. Source: Canned pumpkin. Habitat: L'nknown. 3. Achromobacter sulfureum Bergey et al. (Bakt. 4, Rubentschick, Cent. f. Bakt., II Abt., 72, 1927, 123; Bergey et al., Manual. 3rd ed., 1930, 220.) Rods: 0.7 to 0.8 by 1.7 to 2.2 microns, occurring singlj' and in pairs. Motile. Gram-positive. Gelatin stab: Saccate liquefaction. Agar colonies : Circular, grayish -white, flat, homogeneous. Agar slant: Filiform, grayish -white, smooth, homogeneous growth. Metallic luster. 610 MANUAL OF DETERMINATIVE BACTERIOLOGY Broth: Turbid. Litmus milk: Peptonized. Potato: Yellowish -brown layer. Indole not formed. Nitrites produced from nitrates with gas formation. Hydrogen sulfide formed. Ammonia formed. Urea is attacked. Methylene blue reduced. Aerobic, facultative. Optimum temperature 30° to 33°C. Can grow at 0°C. Source: Sewage filter beds. Habitat: Putrefying materials. Note : See Pseudomonas urease Bergey et al. for another motile, Gram-positive organism described by Rubentschick (Bakt. 3) from the same source. 4. Achromobacter aerophilum (Rubent- schick) Bergey et al. (Urobacterium aerophilum Rubentschick, Cent, f . Bakt., II Abt., 64, 1925, 168; Bergey et al., Manual, 3rd ed., 1930, 224.) Rods : 0.75 to 0.85 by 2.0 to 4.5 microns, occurring singly, in pairs, and in chains. Motile. Gram-positive. Urea gelatin colonies: Small, circular, dirty -gray, entire. Urea gelatin stab: No liquefaction. Urea agar colonies: Circular, grayish, smooth. Urea agar slant : Dirty-gray, glistening to dry growth. Urea broth: Turbid. Urea milk: Unchanged. Urea potato: Slight, grayish-white streak. Indole not formed. Nitrites produced from nitrates. H2S not formed. Ammonia not formed. Aerobic, facultative. Source: Sewage slime. Habitat: Putrefying materials. 5. Achromobacter citrophilum (Ru- bentschick) Bergey et al. (Urobacterium citrophilumRubentschiek, Cent, f . Bakt., II Abt., 64, 1925, 168; 66, 1926, 161; 67, 1926, 167; 68, 1926, 327; Bergey et al., Manual, 3rd ed., 1930, 224.) Rods : 0.75 to 0.85 by 2.5 to 6.0 microns, occurring singly and in pairs. Motile. Gram-positive. Urea gelatin colonies: Small, grayish- white, smooth, undulate. Urea gelatin stab: No liquefaction. Urea agar slant: Filiform, grayish- white, thin, dry growth. Urea broth: Turbid. Urea milk: Unchanged. Urea potato: Dirty-gray, thin streak. Indole not formed. Nitrites produced from nitrates. Hydrogen sulfide not formed. Ammonia not formed. Can derive oxygen from sodium ci- trate. Aerobic, facultative. Optimum temperature 30°C. Source: Sewage slime. Habitat: Putrefying materials. 6. Flavobacterium sulfureum Bergey et al. {Bacterium punctans suljureum Zettnow, Cent. f. Bakt., I Abt., Orig., 77, 1916, 222; Bergey et al., Manual, 1st ed., 1923, 103.) Rods: 0.5 to 0.7 by 0.7 to 1.5 microns. Motile, possessing peritrichous flagella. Gram-positive. Gelatin colonies: Very small, barely visible, becoming brownish -yellow, granular. Gelatin stab: Spreading growth on the surface only. Later crateriform lique- faction. Agar slant: Sulfur-yellow growth. Broth: Turbid. Litmus milk: Alkaline, peptonized, yellow. Potato: Sulfur-yellow streak. Indole not formed. Nitrites not produced from nitrates. Blood serum: Sulfur-yellow growth. Partial liquefaction. Xo acid from glucose. Aerobic, facultative. Optimum temperature 25°C. FAMILY BACTERIACEAE 611 Source: Air. Habitat: Unknown. 7. Flavobacteriinn acetylicum Levine and Soppeland. (Bull. 77, Engineering Exp. Sta., Iowa State Agricultural Col- lege, 1926, 46.) From the chemical term acetyl. Rods: 0.9 by 1.1 microns, with rounded ends, occurring singly and in pairs. Mo- tile. Gram-positive. Gelatin stab: Stratiform liquefaction. Agar colonies: Irregular in form, yel- low, smooth, flat, amorphous, entire. Agar slant: Abundant, echinulate growth, flat, peach yellow, smooth and butyrous. Broth: Ring growth on surface. Tur- bid with scant sediment. Litmus milk: Slight acidity, with granular curd. Pej^tonization. Litmus reduced. Potato: Moderate, orange growth. Indole not formed. Nitrites not produced from nitrates. Starch hydrolyzed. Blood serum liquefied. Acid from glucose with formation of acetylmethylcarbinol . Aerobic, facultative. Optimum temperature 22 °C. Source: From skimmed milk. Habitat: Unknown. 8. Flavobacterium fuscum (Zimmer- mann) Bergey et al. (Bacillus fuscus Zimmermann, Bakt. unserer Trink- und Nutzwasser, Chemnitz, /, 1890, 70; not Bacillus fuscus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 290; Bacterium fuscus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 111; Bergey et al., Manual, 1st ed., 1923, 113; Chromobac- teriurn fuscum Topley and Wilson, Princ. Bact. and Immun., 1, 1931, 405.) From Latin /?/scws, tawny. Rods: 0.6 by 1.5 microns, occurring singly. Xon -motile. Gram-positive. Gelatin colonies: Small, with brownish center and yellowish border. Gelatin stab: Gray, filiform growth in stab. Slow crateriform liquefaction. Agar colonies: Circular, pale yellow, smooth, slightly convex, entire. Agar slant: Growth greenish-yellow, plumose, smooth, raised, undulate. Broth: Turbid, with pellicle and sedi- ment . Litmus milk: Slightly acid, becoming alkaline, with yellow ring. Potato: Thick, moist, chrome-yellow streak. Indole not formed. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 30°C. Source: From Zwonitz River water. Habitat: Water. 8a. Bacterium fuscum liquefuciens (Dyar) Chester. {Bacillus fuscus lique- faciens Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 375; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 108.) Received from the Krai collection labeled Bacillus fuscus; also from air. Differs from the above only in liquefying gelatin more slowly and completely. 9. Flavobacterium maris Harrison. (Canadian Jour. Research, 1, 1929, 232.) From Latin mare, sea. Rods: 0.7 to 0.8 by 1.0 to 1.2 microns, occurring singly and in pairs. At 37°C coccoid. Non-motile. Encapsulated. Gram-positive. Gelatin colonies: Punctiform, red- orange, granular, entire. Gelatin stab: Red-orange surface growth, filiform growth in stab. No liquefaction. Agar colonies : Circular, orange-yellow, smooth, glistening, convex. Agar slant : Growth moderate, orange- yellow, becoming cadmium-orange to red-orange, spreading, glistening. Broth: Clear with orange pellicle and sediment. Litmus milk: At first faintly alkaline, becoming faintly acid with orange sedi- ment. Potato: Scant growth. 612 MAXUAL OF DETERMINATIVE BACTERIOLOGY Indole not formed. Nitrites produced from nitrates. Traces of ammonia formed. Faint acidity from glucose . No action on lactose or sucrose. Loeffier's blood serum not liquefied. No H2S formed. Aerobic, facultative. Optimum temperature 20° to 25°C. Source : Isolated from the skin of fishes. Steinhaus (Jour. Bact., 4^, 1941, 771) found a similar organism in the intestine of a caterisillar. Habitat: Unknown. Appendix II: Among the following species reported as Gram-positive are many that appear to be similar to the species listed at)Ove, while others may belong to the so-called parasitic lacto- bacilli of the intestine or even to Cory- nebacterium. The majority have been placed by their authors in genera other than Bacterium. Bacillus achrous Migula. (No. 6, Lembke, Arch. f. Hyg., 26, 1896, 301; Migula, Syst. d. Bakt., 2, 1900, 676.) From feces. Motile. Bacillus aster if or mis Aligula. (Mi- crobe asteriforme, de Klecki, Ann. Inst. Past., 9, 1895, 735; Migula, Syst. d. Bakt., 2, 1900, 816.) From feces. Motile. Colonies resemble those of Bacillus mycoidcs but no spores are mentioned. Bacillus casei limburgensis Orla-Jen- sen. (Orla- Jensen, Cent. f. Bakt., II Abt., 13, 1904, 702; Bacterium casei lim- burgensis Yale, N. Y. S. Agr. Exp. Sta., Tech. Bui. 268, 1943, 5.) From limbur- ger cheese. Presumably the same as Bacterium linens Weigmann. Bacillus choukevitchi Herter. (Mi- crobe VIII, Choukevitch, Ann. Inst. Past., .^5, 1911, 265; Ilerter, Just's Botan. Jahresber., 39, 2 Abt., Heft 4, 1915, 748.) From the large intestine of a horse. Bacillus colorabilis Migula. (Bacillus der Gallenblase, Nauuyn, Deutsche nied. Wochnschr., 1891, 193; Bacillus cuniculi- cida havaniensis Sternberg, Man. of Bact., 1893, 450; Bacillus coli colorabilis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 434; Bacterium coli colorabilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 87; Bacterium cuniculicida havaniensis Chester, idem; Migula, Syst. d. Bakt., 2, 1900, 736.) From a diseased gall-bladder (Naunyn) and from the intestines and liver of yellow-fever cadavers (Sternberg). Bacillus cupularis Migula. (No. 12, Lembke, Arch. f. Hyg., 26, 1896, 307; Migula, Syst. d. Bakt., 2, 1900, 677.) From feces. Motile. Bacillus fumigatus Migula. (No. 14, Lembke, Arch. f. Hyg., 29, 1897, 320; Migula, Syst. d. Bakt., 2, 1900, 679.) From feces. Motile. Bacillus gazogenes Heurlin. (Heurlin, Bakt. Unters. d. Keimgohaltes im Geni- tal kanale der fiebernden Wochnerinnen. Helsingfors, 1910. 97.) From the vagina. Bacillus glacialis Migula. (Giftpro- duzierender Bacillus, Vaughan and Per- kins, Arch, f . Hyg., 27, 1896, 308; Migula, Syst. d. Bakt., 2, 1900, 807.) From ice cream and cheese. Motile. Bacillus involulus Walsch. (Cent. f. Bakt., I Abt., Orig., 38, 1905, 645.) From preputial secretion. Bacillus laseri Migula. (Gasbildender aerober Bacillus, Laser, Cent., f. Bakt., 13, 1893, 217; Migula, Syst. d. Bakt., 2, 1900, 763.) From the lungs and liver of a calf. Motile. Bacillus membranifer Migula. (No. 7, Lembke, Arch. f. Hyg., 26, 1896, 302; Migula, Syst. d. Bakt., 2, 1900, 675.) From feces. Motile. Bacillus pseudotyphosus Migula. (Ty- phusahnlicher Bacillus, Lustig, Diag. d. Bakt. d. Wassers, 1893, 18; Migula, Syst. d. Bakt., 2, 1900, 730; not Bacillus pseu- dotyphosus Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 383.) Mo- tile. From several samples of water from Thales de Aosta, Italy. Bacillus pulmonum Migula. (Proteus bei Lungengangran des Menschen, Babes, Progres medical roumain, April 6, 1889; Migula, Syst. d. Bakt., 2. 1900, FAMILY BACTERIACEAE 613 755.) From gangrenous lung exudate and the spleen. Motile. Bacillus radiatus Zimmermann. (Zim- mermann, Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, 1, 1890, 58; not Ba- cillus radiatus Liideritz, Ztschr. f. Hyg., 5, 1889, 149; not Bacillus radiatus Ches- ter, Man. Determ. Bact., 1901, 241; Bacillus aquatilis radiatus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 315; quoted as B. radiatus aquatilis Zimmermann by Kruse, idem; Bacterium radiatus aquatilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9; 1897, 100; Bacterium aquatilis radiatus Chester, idem; Bacillus pseudoradiatus IMigula, Syst. d. Bakt., 2, 1900, 830; Bacterium, aquatilis Chester, Man. Determ. Bact., 1901, 160; not Bacterium, aquatile Migula, loc. cit., 469.) From water. Motile. In certain stages of development (pre- sumably old cultures) Zimmermann states that this organism is Gram- negative. Bacillus splendens Migula. {Bacillus ■putidus splendens Bernabei, Bull. d. Soc. Lancisiana d. Ospedali di Roma, 13; see abst. in Cent. f. Bakt., 17, 1895, 469; Migula, Syst. d. Bakt., 2, 1900, 754.) From a case of fetid bronchitis. Motile. Bacterium acidiformans (Sternberg) Chester. (Bacillus acidiformans Stern- berg, Man. of Bact., 1893, 449; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 79.) From the liver of a yellow fever patient. Also reported from bees- wax (White, r. S. Dept. Agr. Bur. En- tomol.. Bull. 14, 1906, 14). Bacterium cactivorum Pasmetti and Buz zati -Traverse. (Xuovo Gior. Bot. Ital., N.S. 42, 1935, 117.) From a dis- eased cactus. Bacterium dendrobii Pavarino. (Rev. di Pat. Veg.. 5, 1912, 242.) From Dend- rohium sp. Bacterium (?) musae (Gaumann) El- liott. (Pseudomonas musae Gaumann, Med. van het Inst, voor Plantz., No. 48, 1921, 58; Elliott, Manual Bact. Plant Path., 1930, 170; Phytomonas (?) m.usae Magrou, in Hauduroy et al.. Diet. d. Bact. Path., 1937, 385.) From diseased bananas. Bacterium nicotinovorum Bucherer. (Cent. f. Bakt., II Abt., 105, 1942, 169.) From a mixture of soil, manure and mud. Bacterium pseudotyphosum ]\Iigula. (Typhusahnlicher Bacillus, Lustig, Diagnostik der Bakterien des Wassers, 1893, 17; Migula, Syst. d. Bakt., 2, 1900, 428.) Non-motile. Isolated by Sartori from Rome tap water. Bacterium teutlium Metcalf. (Met- calf. Cent. f. Bakt., II Abt., 13, 1904, 28; Aplanohacter teutlium E. F. Smith, In- tro, to Bact. Dis. of Plants, 1920, 474; Phytomonas teutlia Bergey et al.. Man- ual, 3rd ed., 1930, 280.) From a soft rot of sugar beets. See Manual, 5th ed., 1939, 216 for a description of this species. Flavobacterium dehydrogenans Ar- naudi. (Micrococcus dehydrogenans Ar- naudi. Boll. Sez. ital. Soc. intern. Micro- biol., 18, 1939, 000; Arnaudi, Cent. f. Bakt., II Abt., 103, 1942-43, 352.) From a pressed yeast cake. Flavobacterium vitarumen Knutsen. (Knutsen, in Bechdel, Honeywell, Dutcher and Knutsen, Jour. Biol. Chem., 80, 1928, 234.) From the rumen of a cow. Kurthia bessoni Severi. (Quoted from Giorn. Batteriol. e Immunol., 34, 1946, 107.) Kurthia variabilis Severi. (Giorn. Batteriol. e Immunol., 34, 1946, 107.) From feces. 22. Bacterium idoneimi McBeth. (McBeth, Soil Science, 1, 1916, 460; Ccllulomonas idonea Bergey et al.. Man- ual, 1st ed., 1923, 165.) From Latin idoneus, capable. Ptods: 0.5 by 1.5 microns. Non -mo- tile. Gram-negative. Gelatin stab: ^Moderate, yellowish growth. Slight napiform liquefaction. Agar colonies: Circular, convex, soft, becoming brittle, grajdsh, granular, en- tire. Agar slant: Scant, yellowish-white growth, becoming distinctly yellow. 614 MANUAL OF DETERMIXATIVE BACTERIOLOGY Gelatin stab: Moderate, yellowish growth. Slight napiform liquefaction. Agar colonies: Circular, convex, soft, becoming brittle, grayish, granular, en- tire. Agar slant: Scant, yellowish-white growth, becoming distinctly yellow. Ammonia cellulose agar: Enzymatic zone shows a diameter of 2 to 3 mm at the end of 30 days. Peptone cellulose agar: Enzymatic zone shows a diameter of 1.5 to 2.0 mm at the end of 30 days. Broth: Turbid. Filter paper broth : Paper reduced to thin, limp sheet which falls apart on slight agitation at end of 15 days. Litmus milk: Acid, not digested. Potato: Abundant, moist, glistening, grayish -white growth, becoming dis- tinctly yellow. Indole not formed. Nitrites produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, starch and glycerol. Aerobic, facultative. Optimum temperature 2G°C. Source: Soil from California. Habitat: Soil. 23. Bacterium udum Kellerman et al. (Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 514; Cellulomonas uda Bergey et al.. Manual, 1st ed., 1923, 166; Proteus cellulomonas var. Proteus udus Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933, 72.) From Latin mlus grow- ing in marshy ground. Rods: 0.5 by 1.5 microns. Non -mo- tile. Gram-negative. Gelatin stab: Liquefaction. Agar slant: Luxuriant, faintly yel- lowish growth. Cellulose agar: Enzymatic zone 0.5 mm wide. Broth: Turbid. Litmus milk: Acid. Potato: Good growth. Indole not formed. Nitrites produced from nitrates. Ammonia is produced. Acid from glucose, fructose, arabinose, xylose, maltose, lactose, sucrose, dextrin and starch. Aerobic, facultative. Optimum temperature 20°C. Source: Compost from Arlington, Va. Habitat: Soil. 24. Bacterium lucrosum McBeth. (McBeth, Soil Science, 1, 1916, 461; Cellulomonas lucrosa Bergey et al., Manual, 1st ed., 1923, 167.) From Latin lucrosus, lucrative. Rods: 0.4 by 1.3 microns. Non-mo- tile. Gram-negative. Gelatin stab: No growth. Agar colonics: Circular, convex, semi- transparent, granular, entire. Aga,r slant: Moderate, flat, grajash- white growth, becoming somewhat iri- descent. Ammonia cellulose agar: On crowded plate, the colonies show an enzymatic zone of 1 mm or more. Peptone cellulose agar: Enzymatic zone 2 to 3 mm wide in 25 days. Broth: Turbid. Filter paper broth: Paper is reduced to a grayish-white pulpy mass whose fibers separate on slight agitation. Litmus milk: No change. Potato: No growth. Indole not formed. Nitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 25. Bacterium acidulimi Kellerman et al. (Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 513; Cellulomonas acidula Bergey et al.. Manual, 1st ed., 1923, 167.) From Latin acidus, acid. Rods : 0.3 by 1 .0 micron. Non-motile. Gram-negative. FAMILY BACTERlACEAE 615 Gelatin stab: No liquefaction. Agar slant: Slight, grayish growth. Cellulose agar: Enzymatic zone 0.5 to 1 mm in width. Broth: Clear. Litmus milk: Acid. Potato: Xo growth. Indole not formed. Nitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose and sucrose. None from gh^cerol or mannitol. Starch not hydroh'zed. Aerobic. Optimum temperature 20°C. Source: Soil from Utah. Habitat: Soil. 26. Bacterium castigatum ^IcBeth. (McBeth, Soil Science, 1, 1916, 458; Cellulomonas casligata Bergey et al., Manual, 1st ed., 1923, 168.) From Latin castigatus, subdued. Rods: 0.4 by 1.2 microns. Non-mo- tile. Gram-negative. Gelatin stab: ^loderate surface growth. No liquefaction. Agar colonies: Circular, slightly con- vex, brittle, grajish-white, granular, entire. Agar slant: Abundant, glistening, grayish-white growth. Ammonia cellulose agar: Enzymatic zone may attain a diameter of 2.5 mm in 30 days. Peptone cellulose agar: Enzj'matic zone may reach a diameter of 2 mm in 30 days. Broth: Slightly turbid. Filter paper broth: Paper completely disintegrated and reduced to a pulp-like mass in 15 da vs. Litmus milk: Acid, not digested. Indole not formed. Nitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch and gh'cerol. Aerobic. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 27. Bacteriimi bibulum (McBeth and Scales) Holland. {Bacillus bibulus'Mc- Beth and Scales, Bur. of Plant Industrj-, U. S. Dept. of Agr., Bui. No. 266, 1913, 35; Holland, Jour. Bact., 5, 1920, 217 and 221; Cellulomonas bibula Berge}' et al., Manual, 1st ed., 1923, 158.) From Latin bibulus, thirsty. Rods: 0.4 by 1.3 microns. ^Motile. Gram-negative. Gelatin stab : Crateriform liquefaction. Cellulose agar colonies: Circular, con- vex, smooth, soft, grayish to faintly yellowish-white, finely granular. En- zymatic zone 0.3 mm in some cases. Agar slant: Luxuriant, glistening, smooth, moist, raised growth. Broth: Slightly turbid. Litmus milk: Faintly acid. Potato: Smooth, glistening, canary yellow growth. Indole is formed. Nitrites not produced from nitrates. Ammonia is produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: From sewer slimes and culti- vated soils. Habitat: Soil. Appendix I : The genus Cellulomonas as originally proposed was based on a single physiological property and included such diverse types of bacteria as (1) polar flagellate species, now placed in Pseudomonas, (2) Gram-variable, non-motile rods now placed in Corynebacterium, and (3) peritrichous, non-spore-forming, Gram-nega- tive rods. Unfortunately the name is unsuitable for the third of these groups so that it has not been inserted in the outline used in this edition of the Manu.a.l. De- scriptions of species previously placed in this genus are given below. 616 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus A. Cellulomonas Bergey el al. (Manual, 1st ed., 1923, 154.) Small rods, with rounded ends, non-spore-forming, motile with peritrichous fla- gella, occurring in soil and having the property of digesting cellulose. Growth on ordinary culture media often not vigorous. Gram-negative. The type species is Cellulomonas biazotea (Kellerman) Bergey et al. Key to the species of genus Cellulomonas. I. Motile with peritrichous flagella. A. Gelatin liquefied. Chromogenic. 1. Milk acid. a. Ammonia not produced; indole not formed. 1. Cellulomonas biazotea. 2. Milk acid; digested. a. Ammonia produced; indole not formed. 2. Cellulomonas aurogenes. aa. Ammonia produced; indole formed. 3. Cellulomonas galba. 3. Milk alkaline. a. Ammonia produced; indole not formed. 4. Cellulomonas folia. 4. Litmus milk unchanged. a. Ammonia produced; indole not formed. 5. Cellulomonas flava. B. Gelatin liquefied. Non-chromogenic. 1. Milk acid. a. Ammonia not produced; indole not formed. 6. Cellulomonas cellasea. aa. Ammonia produced; indole not formed. 7. Cellulomonas iiigis. aaa. Ammonia not produced; indole formed. S. Cellulomonas concitala. 2. Milk acid; digested. a. Ammonia produced; indole not formed. 9. Cellulomonas caesia. C. Gelatin not liquefied. Chromogenic. 1. Milk acid. a. Ammonia produced; indole formed. 10. Cellulomonas gilva. 2. Milk alkaline. a. Ammonia not produced; indole not formed. 11. Cellulomonas ferruginea. D. Gelatin not liquefied. Non-chromogenic. 1. Milk acid; not digested. a. Ammonia not produced; indole not formed. 12. Cellulomonas albida. 13. Cellulomonas alma. aa. Ammonia not produced; indole formed. 14. Cellulomonas desidiosa. aaa. Ammonia produced; indole not formed. FAMILY BACTERIACEAE 617 15. Cellulomonas pusilla. 16. Cellulomonas gclida. II. Motility not recorded. A. Gelatin liquefied. Chroniogenic. 1. Milk acid. a. Ammonia not produced. Acid from glucose. 17. Cellulomonas flavigenu. aa. Ammonia produced. Xo acid from carbohydrates. 18. Cellulomonas rossica. 1. Cellulomonas biazotea (Kellerman et al.) Bergey ot ;i]. (Bacillus biazoteus Kellerman, McBetli, Scales and Smith, Cent. f. Bakt., II Aht.. 39, 1913, 506; Bergey et al., Manual, 1st ed., 1923, 158; Proteus cellulomonas var. Proteus biazoteus Pribram, Klassifikation der Schizomvceten, Leipzig und Wien, 1933, 72.) This is the type species of the genus C ellulomonas . Rods: 0.5 by 0.8 micron. Motile with one to three peritrichous flagella. G ram -nega t i ve . Gelatin stab: Liquefaction. • Agar slant: Luxuriant yellow growth. Cellulose agar: Enzymatic zone 0.25 mm or less in width. Peptone cellulose agar: Xo enzymatic zone. Broth: Turbid. Litmus milk: Acid. Xo curdling or digestion. Potato: Grovv.s well. Indole not formed. X'itrites produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, and glycerol. Xo acid from mannitol. Aerobic, facultative. Optimum temperature 20^C. Source: Soil from Utah. Habitat: Soil. 2. Cellulomonas aurogenes (Kcllernuui et al.) Bergey et al. [Bacillus aurogenes Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 505; Bergey et al.. Manual, 1st ed., 1923, 157.) From Latin and Greek, gold-pro- ducing. Rods: 0.4 by 1.4 microns. Motile with one to three peritrichous flagella. Gram-negative. Gelatin stab: Liquefaction. Agar slant: Luxuriant yellow growth. Cellulose agar: Enzymatic zone 0.5 to 1.5 mm wide. Broth: Turbid. Litmus milk: Acid, digested. Potato: Luxuriant growth. Indole not formed. Xitrites produced from nitrates. Ammonia produced. Acid from glucose, maltose, lactose, sucrose, starch and glycerol. No acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: From soil from Louisiana and Maine. Habitat: Soil. 3. Cellulomonas galba (Kellerman et al.) Bergey et al. [Bacillus galbus Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 509; Bergey et al.. Manual, 1st ed., 1923, 157.) From Latin galbus, yellow. Rods: 0.4 by 1.0 micron. Motile with one to three peritrichous flagella. Gram -negative. Gelatin stab: Liquefaction. Agar slant: Lu.xuriant yellow growth. Cellulose agar: Enzymatic zone 0.5 mm in width. Broth: Turbid. Litmus milk: Acid, digested. Potato: X'o growth. Indole is formed. 618 MANUAL OF DETERMINATIVE BACTERIOLOGY Nitrites not produced from nitrates. Ammonia produced. Acid from glucose, maltose, lactose, sucrose, starch and glycerol. No acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from Louisiana. Habitat: Soil. 4. Cellulomonas folia Sanborn. (Jour. Bact., 1£, 1926, 1 and 343.) From Latin folium, leaf. Description from Sanborn (Jour. Bact., 18, 1929, 170) and also from his unpublished notes. Rods: 0.8 to LO by 1.0 to L5 microns, occurring singly and in short chains. Motile with four to six peritrichous flagella. Gram-negative. Gelatin stab: Slow crateriform lique- faction, becoming stratiform. Agar slant: Growth moderate, dirty- white, echinulate, raised, glistening, opaque, butyrous. Broth: Turbid with yellowish sedi- ment . Litmus milk: Alkaline. Potato: Thick, moist, yellowish-brown growth . Indole not formed. Nitrites produced from nitrates. Acid and gas slowly produced from glucose, sucrose, glycerol and mannitol after prolonged incubation. No acid or gas from lactose. Starch hydrolj^zed. Ammonia produced. No HoS formed. Aerobic, facultative. Optimum temperature 25° to 30°C. Resembles Cellulomonas rossica. .Source: From decomposing leaves. Habitat: Occurring in soil and active in decomposing leaves in composts, hav- ing the property of digesting cellulose. 5. Cellulomonas flava Sack. (Cent. f. Bakt., II Abt., 62, 1924, 79.) From Latin flavus, yellow. Rods: 0.2 by 1.5 microns. Motile. Gram-negative. Gelatin colonies: Circular, citron yellow. Gelatin stab: Very slow liquefaction. Agar colonies: Large, circular, citron yellow. Agar slant: Abundaiat, citron yellow streak. Broth: Turbid with pellicle and sedi- ment. Litmus milk: Unchanged. Potato: Light brown streak. Indole not formed. Nitrites and ammonia produced from nitrates. Hydrogen sulfide produced. Cellulose hydrolyzed. Aerobic, facultative. Optimum temperature 20°C. Habitat: Soil. 6. Cellulomonas cellasea (Kellerman et al.) Bergey et al. {Bacillus cellaseus Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 508; Bergey et al., Manual, 1st ed., 1923, 158.) Rods: 0.5 by 1.2 microns. Motile with one to three peritrichous flagella. Gram-negative. Gelatin stab: Liquefaction. Agar slant: Limited grayish growth. Cellulose agar : Enzj^matic zone 0.5 mm or less. Broth: Clear. Litmus milk: Acid. Potato: No growth. Indole not formed. Nitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from Utah. Habitat: Soil. 7. Cellulomonas iugis (McBeth) Ber- gey et al. (Bacillus iugis McBeth, Soil Science, 1, 1916, 456; Bergey et al.. Manual, Isted., 1923, 158.) From Latin, joined together. Rods: 0.4 by 1.4 microns. Motile with FAMILY BACTEKIACEAE 619 one to three peritrichous flagelki. Gram-negative. Gelatin stab: Xapiform liquefaction. Agar colonies: Circular, convex, soft, grayish-white, granular, entire. Agar slant: Scant, grayish-white, filiform growth. Ammonia cellulose agar: After 20 days, all colonies show an enzymatic zone of 1 mm or more. Peptone cellulose agar: Enzymatic zone continues to increase up to 30 da3'S at which time it may reach 5 mm in width. Broth: Turbid. Filter paper broth: After 15 days, the paper shows many ragged holes but dis- integrates readily. Litmus milk: Acid, not digested. Potato: Abundant, glistening, gray- ish-white growth. Indole not formed. Nitrites produced from nitrates. Ammonia produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 8. Cellulomonas concitata (McBeth) Bergey et al. {Bacillus concitatus Mc- Beth, Soil Science, 1, 1916, 448; Bergey et al.. Manual, 1st ed., 1923, 159.) From Latin cuncitatus, rapid. Rods: 0.5 by 1.2 microns. Motile with one to four peritrichous flagella. Gram-negative. Gelatin stab: Xapiform liquefaction. Agar colonies: Irregularly circular, decidedly convex, soft, becoming viscid, grayish-white, sometimes slightly fluo- rescent, granular, entire. Agar slant: Abundant, flat, moist, faint yellowish-white growth. Ammonia cellulose agar: Surface col- onies show an enzymatic zone of 1.0 to 1.5 mm. Deep colonies no zone but colony somewhat clearer than surround- ing medium. Peptone cellulose agar: Lnzjiuatic zone, surface colonies, 2 to 2.5 mm; bot- tom colonies, 1 mm or less. Broth: Turbid. Filter paper broth; In 15 days, the paper is a disintegrated fibrous mass which retains its pure white color. Litmus milk: Acid, not digested. Potato: No growth. Indole is formed. Nitrites not produced from nitrates. Ammonia not produced. Acid from fructose, maltose, lactose, sucrose, starch and glycerol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 9. Cellulomonas caesia (Kellerman et al.) Bergey et al. {Bacillus caesius Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 507; Cellulomonas caseia (sic) Bergey et al., Manual, 1st ed., 1923, 159; Cellulomonas casei (sic) Bergey et al., Manual, 4th ed., 1934, 199.) From Latin caesius, bluish- gray. Rods: 0.4 by 1.5 microns. Motile with one or two peritrichous flagella. Gram- negative. Gelatin stab: Liquefaction. Beef agar streak: Moderate, flat, thin growth, slightly bluish fluorescence. Cellulose agar: Enz^-matic zone, 0.5 to 1.0 mm in 15 days. Broth: Turbid. Slight sediment in 5 days. Litmus milk: Acid, digested. Potato: No growth. Indole not formed. Nitrites produced from nitrates. Ammonia produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from Louisiana, Wiscon- sin and New Hampshire. Habitat: Soil. 620 MANUAL OF DETERMINATIVE BACTERIOLOGY 10. Cellulomonas gilva (McBeth) Ber- gey et al. {Bacillus gilvus AIcBeth, Soil Science, 1, 1916, 453; Bergey et al.. Manual, 1st ed., 1923, 160.) From Latin gilvus, pale yellow. Rods: 0.5 by 1.5 microns. Motile with one to five peritrichous flagella. Gram-negative. Gelatin stab: Moderate, yellowish- white surface growth. Xo liquefaction. Agar colonies: Circular, convex, bu- tyrous, canary -yellow, sometimes with brownish rings, granular, entire. Agar slant: Filiform, yellowish-white growth. Ammonia cellulose agar: Enzj-mic zone not more than 1 mm. Entire col- ony semi transparent. Peptone cellulose agar: Enzymatic zone, 3 to 4 mm in 25 days. Broth: Slightly turbid. Filter paper broth: In 15 days, the paper is reduced to a thin, white filmy mass which disintegrates readily. Litmus milk: Acid, not digested. Potato: Abundant, canary -yellow growth . Indole is formed. Nitrites produced from nitrates. Ammonia is produced. Acid from glucose, maltose, lactose, sucrose, starch and glj'cerol. Xo acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 11. Cellulomonas ferruginea (Uull- mann) Bergey et al. (Bacillus ferru- gineus Ilullmann, Cent. f. Bakt., I Abt., Orig., 24, 1898, 465; van Iterson, Cent, f. Bakt., II Abt., 11, 1904, 694; not Bacillus ferrugineus Rullmann, Cent. f. Bakt., I Abt., 24, 1898, 467; Bergey ct al., Manual, 1st ed., 1923, 150.) From Latin, rust-colored. Rods: 0.5 to 0.8 by 1.5 to 2.0 microns, occurring singly. Motile, possessing peritrichous flagella. Gram-negative. Gelatin colonies: Brown, the pigment dift'using into the medium. Gelatin stab: Xo liquefaction. Agar slant: Rusty-brown streak. Broth: Turbid. Litmus milk: Dark-yellow ring; alka- line. Potato: Rusty-brown streak. Indole not formed. Xitrites not produced from nitrates. Ammonia not produced. Aerobic, facultative. Optimum temperature 25°C. Habitat: Water. 12. Cellulomonas albida (McBeth) Ber- gey et al. {Bacillus albidus McBeth, Soil Science, 1, 1916, 445; Bergey et al., Manual, 1st ed., 1923, 160.) From Latin albidus, white. Rods-. 0.4 by 1 .0 micron. Motile with one to three peritrichous flagella. Gram-negative. Gelatin stab : Scant growth . Xo lique- faction. Agar colonies: Circular, convex, soft, grayish -white, granular, entire. Agar slant: Scant, white streak. Ammonia cellulose agar : After 30 days the colonies show an enzymatic zone of 1 to 2 mm. Broth: Clear. Filter paper broth: In" 15 days, the paper is reduced to a thin, filmy, grayish- white mass which readilj' breaks up. Litmus milk: Slightly acid, not di- gested. Potato: Xo growth. Indole not formed. Xitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch, glycerol and mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 13. Cellulomonas alma (McBeth) Ber- gey et al. {Bacillus almics McBeth, Soil Science, 1, 1916, 446; Bergey et al., FAMILY BACTERIACEAE 621 Manual, 1st ed., 1923, 161.) From Latin almus, nourishing. Rods : 0.5 by 1.2 microns. Motile with one to five peritrichous flagella. Gram- negative. Gelatin stab : Scant growth. Xo lique- faction. Agar colonies: Circular, convex, soft, becoming brittle, grayish-white, granu- lar, entire. Ammonia cellulose agar: Enzj^matic zone 3 to 4 mm in 25 days. Peptone cellulose agar: Enzymatic zone 2.5 to 3.5 mm in 30 days. Agar slant: Scant, grayish-white growth, becoming yellowish-white. Broth: Slightly turbid. Filter paper broth: Paper reduced to a loose felt -like white mass in 15 days. Litmus milk: Slightly acid, not di- gested. Potato: Xo growth. Indole not formed. Xitrites not produced from nitrates. Ammonia not produced. Acid from glucose, maltose, lactose, sucrose, starch and glycerol. Xo acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 14. Cellulomonas desidiosa (McBeth) Breed. {Bacillus desiduosus (sic) Mc- Beth, Soil Science, 1, 1916, 450; Cellulo- monas deciduosa (sic) Bergey et al., Manual, 1st ed., 1923, 162; Breed, in Manual, 5th ed., 1939, 495.) From Latin desidiosus, inactive. Rods: 0.4 by 1.0 micron. Motile with one to three peritrichous flagella. Gram-negative. Gelatin stab: Moderate growth. Xo liquefaction. Agar colonies: Circular, slightly con- vex, soft, becoming somewhat viscid, grayish-white, granular, entire. Agar slant: Scant, flat, grayish -white growth . Ammonia cellulose agar: Enzymatic zone 3 to 3.5 mm in 25 days. Peptone cellulose agar: Enzymatic zone 1 to 2 mm around surface colonies. Bottom colonies frequently show no en- zymatic zone until after 20 days. Broth: Slightly turbid. Filter paper broth: Paper is divided into gray white mass which readily dis- integrates. Litmus milk: Acid, not digested. Potato: Xo growth. Indole is formed. Xitrites produced from nitrates. Ammonia not produced. Acid from glucose, lactose, maltose and starch. X^o acid from mannitol, sucrose or glycerol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from California. Habitat: Soil. 15. Cellulomonas pusilla (Kellerman et al.) Bergey et al. {Bacillus pusilus (sic) Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 513; Cellulomonas pusila (sic) Bergey et al.. Manual, 1st ed., 1923, 161.) From Latin pusilla, very small. Rods: 0.6 by 1.1 microns. Motile with one to three peritrichous flagella. Gram -negative. Gelatin stab: Xo liquefaction. Agar slant: Scant, grayish -white growth. Cellulose agar: Enzymatic zone i mm or less in width. Broth: Turbid. Litmus milk: Acid. Potato: Xo growth. Indole not formed. Xitrites produced from nitrates. Ammonia is produced. Acid fi'om glucose, maltose, lactose, sucrose, starch and glycerol. Xo acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. 622 MANUAL OF DETERMTN'ATIYE BACTEUIOLOGY Source : Soil from District of Columbia and South Carolina. Habitat: Soil. 16. Cellulomonas gelida (Kellerman et al.) Bergey et al. (Bacillus gelidus Kellerman, McBeth, Scales and Smith, Cent. f. Bakt., II Abt., 39, 1913, 510; Bergey et al., Manual, 1st ed., 1923, 162.) From Latin gelidus, stiff. Rods: 0.4 by 1.2 microns. Motile with one to three peritrichous flagella. Gram -negative. Gelatin stab: No liquefaction. Agar slant: Lu.xuriant, grayish-white growth. Cellulose agar: Enzymatic zone 1.5 mm in width. Broth: Turbid. Litmus milk: Acid, peptonized. Potato: Grows well. Indole not formed. Nitrites not produced from nitrates. Ammonia is produced. Acid from glucose, maltose, lactose, sucrose, starch and glycerol. No acid from mannitol. Aerobic, facultative. Optimum temperature 20°C. Source: Soil from Connecticut. Habitat: Soil. 17. Cellulomonas flavigena (Kellerman and McBeth) Bergey et al. {Bacillus flavigena Kellerman and McBeth, Cent, f. Bakt., II Abt., 34, 1912, 488; Bergey et al., Manual, 1st ed., 1923, 165.) From Latin, yellow-producing. Rods: 0.4 by 1.0 micron. Motility not recorded. Gram-negative. Gelatin stab: Liquefaction. Agar slant: Luxuriant, yellow growth. Cellulose agar: Enzymatic zone 0.75 to 1.5 mm in width. Broth: Turbid. Litmus milk: Acid. Potato: Grows well. Indole not formed. Nitrites produced from nitrates. Ammonia not produced. Acid from glucose, fructose, arabinose, xylose, maltose, lactose, sucrose, dextrin, starch, inulin, salicin, glycerol and man- nitol. Aerobic, facultative. Optimum temperature 20°C. Source: From contaminated culture. Habitat: Soil. 18. Cellulomonas rossica (Kellerman and McBeth) Bergey et al. (Bacillus rossicus Kellerman and McBeth, Cent, f. Bakt., II Abt., 34, 1912, 492; Bergey et al., Manual, 1st ed., 1923, 157; Proteus cellulomonas var. Prote^is rossicus Prib- ram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 72.) Rods: 0.3 by 1.2 microns. Motility not recorded. Gram-negative. Gelatin stab: Rapid liciuefaction. Agar slant: Luxuriant, yellow growth. Cellulose agar: Enzymatic zone 0.5 to 1.0 mm in width. Broth: Turbid. Litmus milk: Alkaline. Potato: Grows well. Indole not formed. Nitrites produced from nitrates. Ammonia produced. No acid from carbohydrate media. Aerobic, facultative. Optimum temperature 20°C. Source: From contaminated culture. Habitat: Soil. Appendix II : Tlie following cellulose- digesting bacteria are not included above: Achromobacler picru7n Fuller and Nor- man. (Jour. Bact., 46, 1943, 276.) From soil. Bacillus aurogenes var. albus Keller- man, McBeth, Scales and Smith. (Cent, f. Bakt., II Abt., 39, 1913, 506.) From soil from New York State. Differs from Cellulomonas aurogenes in that it shows no chromogenesis. Bacillus rossicus var. castaneus Keller- man et al. (loc. cit., 508; Proteus cellu- lomonas var. Proteus castaneus Pribram, Klassifikation der Schizomyceten, Leip- zig und Wien, 1933, 72.) From «oils FAMILY BACTERIACEAE 623 from Maine, Connecticut and New York. Peritrichous. Xo liquefaction of gela- tin. Chestnut color on potato. Bacillus subalbus Kellerman et al. (loc. cit., 512) . From soils from Georgia, Kentucky and New York. Bacillus suhalhus var. batatatis Keller- man etal. (loc. cit., 513). From soil from South Carolina. Differs from the above species in that it liquefies gelatin and forms a very scant yellowish growth on potato. Appendix III : The following genus has been proposed for Gram-negative rods that utilize bacterial polysaccharides as a sole source of carbon. Genus A. Saccharobacterium Sickles and Shaw. (Jour. Bact., 28, 1934, 430.) Pleomorphic, non-motile, non-spore-forming rods. Gram-negative. Grow in mineral solutions containing bacterial polysaccharicks as the sole source of carbon. Found in swamp and other uncultivated soils. Placed by the authors in the Family Mycobactcriaccae because of resemblances between these bacteria and those placed in Cytophaga, Cellfalcicula and Cellvibrio. As the latter genera are no longer placed in this family, Saccharobacterium is placed temporarily in this appendix to the genus Bacterium near bacteria that decompose cellulose and agar. The type species is Saccharobacterium ovale Sickles and Shaw. 1 . Saccharobacterium ovale Sickles and Shaw. (Jour. Bact., 28, 1934, 422.) From Latin ovum, egg, ellipse. Extremely pleomorphic. Young cells ellipsoidal, 1.5 bj' 2.0 microns, usually in pairs, contain granules which stain deeply with basic dyes. Older cultures contain cells which may be from 12 to 15 microns long. Non-motile. Gram- negative. No growth on ordinary media such as beef-extract agar, blood agar, beef-ex- tract agar slants, nutrient gelatin, po- tato slants, litmus milk, beef -infusion broth and beef -extract peptone broth. Medium A plus pneumococcus II car- bohydrate and O.S per cent agar: Very small, round, pink colonies, pinpoint in size after about 5 days. After 2 weeks 1 mm in diameter. Coherent. Litmus milk: Xo growth. Beef-extract peptone with 1 per cent sucrose : Moderate turbidity. Yellowish sediment. Starch : Hydrolyzed in Medium A con- taining pneumococcus II carbohydrate. Growth in lactose and sucrose broths. Growth in maltose, xylose and dextrin broths in some strains. No acid from inulin, mannitol, salicin and glucose broths. Aerobic. Minimum temperature 20°C. Opti- mum 34° to 35 °C. Maximum 37 °C. Thermal death point 54°C for 10 minutes. Minimum pH 6.4. Optimum pH 7.0 to 7.4. Maximum pH 7.8. Distinctive characters: The addition of 0.5 per cent sodium chloride to any favorable medium completely prevents growth of the organism (Medium A is that used by Dubos and Avery in 1931, (NH4)2S04, 1 g, K2HPO4, 2.0 g, tap water 1000 ml) . Decomposes the carbohydrate of pneumococcus type II. Source: Swamps and other unculti- vated soils. Habitat: SoiL 2. Saccharobacteriiun acuminattun Sickles and Shaw. (Jour. Bact., 28, 1934, 425.) From Latin acuminare, to sharpen. Extremely pleomorphic. Young or- ganisms are pointed, often curved rods, 624 MANUAL OF DETERMINATIVE BACTERIOLOGY 0.5 by 2 microns, having a densely stain- ing granule. The tapering pointed ends remain unstained. Older cells have rounded ends, are spherical, pear- shaped or a long ellipsoid, stain weakly. Non-motile. Gram-negative. No growth on ordinary media. See preceding species. Medium S with pneumococcus I car- bohydrate and 0.8 per cent agar: Very tiny, pale yellow colonies. Less than 0.5 mm in diameter. Starch not hydrolyzed. Growth in sucrose broth. No growth in glucose, lactose, maltosef dextrin, inulin, mannitol and saliein broths. Aerobic. Minimum temperature 20°C. Opti- mum 28° to 32°C. Maximum 34°C. Thermal death point 48°C for 10 minutes. Minimum pH 6.0. Optimum pH 6.6 to 7.2. Maximum pH 7.8. Distinctive characters: Decomposes the carbohydrate of pneumococcus Type I. The addition to any favorable medium of 0.7 per cent sodium chloride, of 0.3 per cent beef extract or of 0.5 per cent peptone completely inhibits growth. The composition of Medium S is as follows: MgS07H20, 0.2 g, NH4H2PO4, 1.5 g, CaCl2, 0.1 g, FeCh, tr, KCl, 0.1 g, 10 cc N/1 NaOH. Distilled water 1,000 ml, pH 7.2 to 7.4. To this was added the specific pneumococcus carbohydrate as a source of carbon in concentrations varying from 0.002 to 0.01 per cent. Source: From swamps and other un- cultivated soils. Habitat: Soil. 28. Bacterium nenckii Biernacki. (Biernacki, Cent. f. Bakt., II Aht.,.29, 1911, 166; Achromobacter nenckii Bergey et al.. Manual, 3rd ed., 1930, 227.) Named for Nencki, a chemist at the Medical Institute in Warsaw. Rods: 0.8 by 1.25 to 2.0 microns, with rounded ends, occurring singly and in pairs. Capsulated. Non-motile. Gram-negative. Gelatin colonies: Circular, convex, yellowish -white, granular. Glucose and sucrose gelatin: Colonies larger and slimy. Gelatin stab: No liquefaction. Agar colonies : Circular, grayish -white, glistening, concentric, finely granular. Agar slant: The medium is liquefied. Glucose and sucrose agar : Heavy slimy growth with gas. Faint fruity odor. Broth: Slightly turbid with gray sedi- ment and slight odor. Litmus milk: Acid and gas formation. Potato: Slight growth. Glycerol potato: Heavy growth with the appearance and consistency of cream. Indole not formed. Nitrites not produced from nitrates. Acid and gas from glucose, fructose, galactose, maltose, sucrose, raffinose and mannitol. Fruity odor in cultures. Facultative anaerobe. Optimum temperature 37°C. Source: From Spanish dried grapes. Habitat: Unknown. 29. Bacterium polysiphoniae Lunde- stad. (Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 331; Flavobacterium poly- siphoniae Bergey et al., Manual, 3rd ed., 1930, 152.) From Greek, many tubes. Rods: 0.5 to 0.6 by 2.0 to 4.0 microns, with rounded ends, occurring singly. Non-motile. Gram-negative. Fish-gelatin colonies: Circular, slightly glistening, bright yellow, trans- parent, with denser center. Fish-gelatin stab: Slight yellowish growth on surface. Slow saccate lique- faction. Sea-weed agar colonies: Circular, flat, with concentric rings, diffuse margin, light yellow. Agar is disintegrated. Fish-agar slant: Yellow, flat growth, with undulate margin. Broth : Turbid with flocculent pellicle and yellowish sediment. Indole not formed. FAMILY BACTERIACEAE 625 Nitrites not produced from nitrates. No action on carbohydrates. Slight hydrolysis of starch. Aerobic, facultative. Optimum temperature 30°C. Source: Sea water of Norwegian Coast. Habitat: Sea water. 30. Bacterium droebachense Lunde- stad. (Lundestad, Cent. f. Bakt., II Abt., 1928, 329; Flavobacterium droe- bachense Bergey et al., Manual, 3rd ed., 1930, 153; Pseudomonas droebachense Stanier, Jour. Bact., 4^, 1941, 544.) Latinized, from Drobak, where this or- ganism was isolated. Rods: 0.5 to 0.6 by 2.0 to 2.6 microns, with rounded ends, occurring singly. Non-motile. Gram-negative. Fish-gelatin colonies: Small, circular, compact, opaque, glistening, orange- yellow. Fish-gelatin stab: Liquefaction in- fundibuliform becoming stratiform. Sea-weed agar colonies: Small, circu- lar, flat, opaque, glistening, orange- yellow. Agar is disintegrated. Fish-agar slant: Flat, opaque, glisten- ing, slimy, orange-yellow, entire growth. Broth: Slight flocculent turbidity, yellow. Indole not formed. Nitrites not produced from nitrates. Starch hydrolyzed. Slow growth on surface of glucose agar stab. No gas. Aerobic, facultative. Optimum temperature about 37°C. Minimum temperature 5° to 10°C. Maximum 40 °C. Stanier (loc. cit.) identified cultures isolated from sea water on the Pacific Coast as belonging to this species. Some liquefied gelatin while others did not. Nitrates were reduced. A yellow membraneous pellicle was formed on broth, and the temperature range is given as 5° to 35°C. Optimum 25°C. He renamed the organism Pseudomonas droebachensis, but reported it non- motile. Source: From sea water at Drobak on the Norwegian Coast. Habitat: Sea water. 31. Bacterium delesseriae Lundestad. (Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 332; Flavobacterium delesseriae Bergey et al.. Manual, 3rd ed., 1930, 153.) Rods: 0.5 to 0.6 by 1.6 to 2.6 microns, with rounded ends, occurring singly. Non-motile. Gram-negative. Fish-gelatin colonies: Circular, trans- parent, glistening, concentrically ringed, yellow. Fish -gelatin stab: Crateriform lique- faction, with yellow sediment. Sea-weed agar colonies: Circular, flat, concentrically ringed, light yellow. Agar is disintegrated. Fish-agar slant: No growth. Broth : Turbid with flocculent pellicle and sediment, light yellow. Indole not reported. Nitrites not reported. No action on carbohydrates. Slight hj'drolysis of starch. Aerobic, facultative. Optimum temperature 23°C. Source : Sea water of Norwegian Coast. Habitat: Sea water. 32. Bacterium boreale Lundestad. (Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 333; Flavobacterium boreale Bergey et al., Manual, 3rd ed., 1930, 154.) From Latin borealis, northern. Rods: 0.5 to 0.6 by 1.6 to 2.6 microns, with rounded ends, occurring singly. Non-motile. Gram-negative. Fish -gelatin colonies: Circular, opaque, glistening, concentrically ringed, yellow. Fish-gelatin stab : Yellow, with crateri- form liquefaction. Sea-water agar colonies: Circular, flat, opaque, glistening, diffuse margin, light yellow. Agar is disintegrated. 626 MANUAL OF DETERMINATIVE BACTERIOLOGY Fish-agar slant: Yellow, flat, glisten- ing, opaque, entire growth. Broth: Finely flocculent, yellow sedi- ment . Indole not reported. Nitrites not reported. No action on carbohydrates. Slight hydrolysis of starch. Aerobic, facultative. Optimum temperature 23°C. Source : Sea water of Norwegian Coast. Habitat: Sea water. 33. Bacterium ceramicola Lundestad. (Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 332; Flavohacteriian ceramicola Bergey et al., Manual, 3rd ed., 1930, 154.) From Greek, living in earthenware. Rods: 0.5 to 0.6 by 1.4 to 2.4 microns, with rounded ends, occurring singly and lying side-by-side. Non-motile. Gram- negative. Fish-gelatin colonies: Circular, glis- tening, transparent, yellow. Fish-gelatin stab: Slight, yellow sur- face growth. Liquefaction crateriform. Sea-water agar colonies: Circular, flat, transparent, glistening, diffuse margin, light 3'ellow. Agar is disintegrated. Fish-agar slant: Moderate, yellow, flat, entire, glistening, opaque growth. Broth: Light yellow pellicle and sedi- ment. Indole not reported. Nitrites not reported. No action on carbohydrates. Slight hydrolysis of starch. Aerobic, facultative. Optimum temperature 23°C. Source: Sea water of Norwegian Coast. Habitat: Sea water. 34. Bacterium rhodomelae Lundestad. (Lundestad, Cent. f. Bakt., II Abt., 75, 1928, 331; Flavobacterium rhodomelae Bergey et al.. Manual, 3rd ed., 1930, 146.) Rods: 0.5 to 0.8 by 1.2 to 2.0 microns, with rounded ends, occurring singly, in })airs, and at times in short chains. Mo- tile. Gram-negative. Fish-gelatin colonies: Circular, slightly glistening, opaque, white. Fish-gelatin stab: Rapid infundibuli- form liquefaction. Sea-weed agar colonies: Circular, flat, thin, transparent, glistening, entire. Agar is dissolved. Glucose agar slant: Moderate growth, white, becoming orange-yellow, flat, un- dulate margin, opaque, glistening. Broth : Turbid, with pellicle and gray- ish-yellow, slimy sediment. Indole not formed. Nitrites not produced from nitrates. No action on carbohydrates. Very slight hydrolysis of starch. .\erobic, facultative. Optimum temperature 20° to 25°C. Source : Sea water of Norwegian Coast. Habitat: Sea water. 35. Bacteriiun alginovorum Waksman, Carey and Allen. (Jour. Bact., 28, 1934, 215.) From M. L., alginic and Latin voro, devour. Rods: 0.75 to 1.2 by 1.5 to 2.0 microns, with rounded to almost elliptical ends, especially when single, occurring fre- quentl}^ in pairs and even in chains. Actively motile. Capsule-forming. Gram-negative. Alginic acid plate : Colony large, white in appearance with coarse granular cen- ter, entire margin. Clears up turbidity caused by the alginic acid on plate. No odor. Alginic acid liquid medium: Heavy pellicle formation. Active production of an enzyme, alginase, which brings about the disappearance of alginic pre- cipitate in sea water medium. Salt water medium: A slimy pellicle of a highly tenacious nature is produced, the whole medium later turning to a soft jelly. Sea water gelatin: Active and rapid liquefaction in two to six days, at 18°C; FAMILY BACTERIACEAE 627 highly turbid throughout the liquefied zone. Agar liquefaction: Extensive softening of agar, no free liquid. Sea water glucose broth: Abundant uniform turbidity, with surface pellicle; some strains give heavier turbidity and others heavier pellicle. Litmus milk containing 3.5 per cent salt: No apparent growth. Potato moistened with sea water: Moist, spreading growth, ivory-colored; heavy sediment in free liquid at the bottom. Starchplate: Abundant, cream-colored, slimy growth; extensive diastase produc- tion. Aerobic, microaerophilic. Optimum temperature 20°C. Source: From sea water, sea bottom sediments and from the surface of algal growth in the sea. Habitat: Very common in the sea. 36. Bacterium fucicola Waksmau, Carey and Allen. (Jour. Bact., 28, 1934, 213.) From Latin fucus, seaweed and cola, dweller. Short rods: 0.6 to 1.0 bv 1.0 to L5 microns, with ends rounded to almost coccoid; slightly curved. Actively mo- tile, with twirling motion. Gram-nega- tive. Alginic acid plate: Colonies finely granular, entire; at first whitish, turning brown in three to five days, and later almost black, producing a deep brown soluble pigment. Alginic acid liquid medium: Limited growth on surface in the form of a pelli- cle. Frequently produces no growth at all. Sea water gelatin: Active liquefaction; no growth in stab; thin, fluorescent growth throughout liquefied zone. Agar liquefaction: Positive, although limited; only softening of agar. Sea water glucose broth: Faint tur- bidity; no pellicle, no sediment. Litmus milk containing salt: No ap- parent growth. Potato moistened with sea water: Xo growth. Starch plate: Xo growth. Aerobic. Optimum temperature 20°C. Source: From sea water near the sur- face of the sand bottom. Habitat: Rare in sea water. Appendix I: Additional agar-digesting bacteria placed in genera other than Bac- terium. \. Achromobacter. A. Motile with peritrichous flagella. 1. Xitrites produced from nitrates. 1. Achromobacter pastinalar. II. Agarbacterium. A. Xon-motile. 1. Xitrites produced from nitrates. 2. Agarbacterium bufo. B. Motile, but position of flagella not recorded. 1. Xitrites produced from nitrates. 3. Agarbacterium reducans. 4. Agarbacterium viscosuyn. 2. Xitrites not produced from nitrates. 5. Agarbacterium mesentericus. 6. Agarbacterium aurantiacum. 7. Agarbacterium cyanoidcs. 3. Seven additional species that are numbered but not named. 628 MANUAL OF DETERMINATIVE BACTERIOLOGY III. Flavobacterium . A. Non-motile. 1. Nitrites produced from nitrates. 8. Flavohactcrium uliginosum. B. Peritrichous flagella. 1. Nitrites produced from nitrates. 9. Flavobacterium ainocontacturn. 1 . Achromobacter pastinator Goresline. (Jour. Bact., 26, 1933. 442.) From Latin pastinator, one who digs a trench. Short rods: 0.4 by 1.5 microns, occur- ring singly and in pairs. Motile with two to five peritrichous flagella. Gram- negative. Plain gelatin stab: No growth. Nutrient gelatin stab : Surface growth very scanty. No liquefaction. Nutrient agar colonies: At first tiny, almost colorless, becoming yellowish and ring-like. Agar liquefied rapidly. Nutrient agar slant : Growth good, fiat, not thick. Agar liquefied along streak often to the depth of a quarter of an inch. Pocket formed at bottom of slant filled with a rather viscous, yellowish fluid. Nutrient broth: Slight turbidity after 5 days. Subsurface but no surface growth. No sediment. Litmus milk: Slightly acid after 20 days. No curd. Only a trace of reduc- tion at bottom of tube. Potato: No growth. Indole not formed. Nitrites produced from nitrates. No H2S produced. Acid from arabinose, glucose, galac- tose, lactose, fructose, maltose, mannose, melezitose, pectin, raffinose, rhamnose, salicin, sucrose, starch and de.xtrin. No growth in dulcitol, erythritol, manni- tol, sorbitol, glycerol, xylose and inulin. Starch is hydrolyzed. Limits of growth: pH .5.9 to 9.0. Temperature relations: Optimum 28°C. Good growth at 25^0. Moderate growth at 20° and 37°C. No growth at 10° and 42°C. Facultative anaerobe. Distinctive characters: Digests agar rapidly; colonies sink through to the glass of the Petri dish. Fehling's solu- tion reduced by the liquefied agar. Con- siderable change in viscosity of agar due to the digestion. Source: From a trickling filter receiv- ing creamery wastes. Habitat: Probably widely distributed in nature. 2. Agarbacterium bufo Angst. (Puget Sound Biol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.6 by 0.8 micron, occurring singly and in pairs. Non-motile. No capsules. Gram-nega- tive. Fish gelatin stab: Stratiform liquefac- tion, growth best at top. Fish gelatin colonies: Circular, crateri- form, granular. Fish agar slant: Abundant, filiform, raised, glistening, opaque, yellow, mem- branous growth. Fish agar colonies: Circular, concen- trically ringed, sunken, entire, granular, yellow to orange. Digests agar; cellulose not attacked. Potato: No growth. Plain milk unchanged, surface growth yellow. Does not produce H2S or indole. Nitrites produced from nitrates. .Vcid from mannitol. No acid from xylose, rhamnose, arabinose, glucose, sucrose or lactose. Starch not hydrolj'zed. Aerobic. Optimum temperature 25° to 2S°C. Maximum under 36 °C. Source : Isolated from Odonthalia kamt- schatica. Habitat: On marine algae. FAMILY BACTERIACEAE 629 3. Agarbacterium reducans Angst. (Puget Sound Biol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.6 by 0.8 micron, occurring singly and in pairs. Motile. No capsules. Gram-negative. Fish gelatin colonies : Circular, sunken, entire, crateriform, granular. Fish gelatin stab: Crateriform lique- faction, growth only near surface. Fish agar slant: Abundant, filiform, flat, glistening, smooth, opaque, white, butyrous growth. Fish agar colonies: Moderate, circular, smooth, flat, entire, granular, white to buff or colorless. Digests agar; cellulose not attacked. Fish broth: Turbid, no sediment, no surface growth. Potato: No growth. Nitrites produced from nitrates. No H2S or indole formed. Plain milk unchanged. Acid from sucrose, arabinose, rham- nose and mannitol . No acid from .xylose or lactose. Starch is hj'drolyzed. Aerobic. Optimum temperature 25° to 28°C; thermosensitive. Source: Isolated from NercocysLis luetkeana. Habitat: On marine algae. 4. Agarbacterium viscosum Angst. (Pugot Sound P.iol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.6 to 0.8 micron, occurring singlj' or in pairs. Motile. No capsules. Gram-negative. Fish gelatin colonies : Circular, sunken , entire, crateriform, granular. Fish gelatin stab: Stratiforrn liquefac- tion, growth best at surface. Fish agar slant: Abundant, raised, glistening, smooth, opaque, graj% vesicu- lar, viscid growth. Fish agar colonies: Circular, con- toured, raised, lobate, granular, fluores- cent green. Digests agar; cellulose not attacked. Fish broth: Flocculent pellicle, turbid, no sediment, fluorescent green. Potato: Abundant, filiform, glistening, smooth, yellowish-brown, butyrous growth. Nitrites produced from nitrates. No H2S or indole formed. Plain milk unchanged; surface growth greenish. No acid from rhamnose, sucrose, lac- tose, mannitol, xylose or arabinose. Starch is hydrolyzed. Aerobic. Optimum temperature 20° to 28°C; thermosensitive. Source: Isolated from Iridaea cordata. Habitat: On marine algae. 5. Agarbacteriimi mesentericus Angst. (Pugot Sound Biol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.6 by 0.8 micron, occurring singly or in pairs. Motile. No capsules. Grp.m-negative. Fish gelatin stab: Infundibuliform liquefaction; growth best at top. Gelatin colonies: Circular, sunken, irregular, crateriform, granular. Fish agar slant: Abundant, filiform, raised, glistening, finely wrinkled when old or dry, opaque, buff, membranous growth. Fish agar colonies: Circular, concen- trically ringed, flat, entire, granular, white to buff. Digests agar; cellulose not attacked. Fish broth: Membranous pellicle, moderate clouding, no sediment. Potato: Spreading, raised, glistening, wrinkled, buff to yellowish, membranous growth . Does not produce H^S or indole. Nitrites not produced from nitrates. Plain milk unchanged. Acid from mannitol. No acid from xylose, rhamnose, arabinose, glucose or lactose. Starch is hydrolyzed. Aerobic. Optimum temperature 20° to 28°C; thermosensitive . 630 MANUAL OF DETERMINATIVE BACTERIOLOGY Source: Marine algae; isolated from NercGcystis luetkeana. Habitat: On marine algae. 6. Agarbacterium aurantiaciun Angst. (Puget Sound Biol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.6 to 0.8 micron, occurring singly or in pairs. Motile. No capsules. Gram-negative. Fish gelatin colonies : Circular, sunken, crateriform, entire. Fish gelatin stab: Stratiform liquefac- tion, no growth along line of stab. Fish agar slant: Abundant, filiform, flat, glistening, smooth, opaque, orange, butyrous growth. Fish agar colonies: Circular, smooth. Hat, erose, sunken, granular. Digests agar; cellulose not attacked. Fish broth: Membranous pellicle, tur- l)id, no sediment. Plain milk unchanged; surface growth orange. Potato: Alumdant, filiform, flat, dull, smooth, orange, butyrous growth. Nitrites not produced from nitrates. No HoS or indole formed. Acid from lactose and mannitol. No acid from xylose, rhamnose, arabinose, glucose or sucrose. Starch not hydrolyzed. Aerobic. Optimum temperature 20° to 28°C; thermosensitive. Source: Isolated from Porphi/ra per- forata. Habitat: On marine algae. 7. Agarbacterixim cyanoides Angst. (Puget Sound Biol. Sta. Pub., 7, 1929, 49.) Short rods with rounded ends, 0.8 by 1.4 microns, occurring singly or in pairs. Motile. No capsules. Gram-negative. Fish gelatin colonies : Circular, sunken , entire, crateriform, granular. Fish gelatin stab: Stratiform liquefac- tion, growth only at top. Fish agar slant: Abundant, filiform, raised, glistening, smooth, opaque, gray, butyrous growth. Fish agar colonies: Circular, smooth, flat, lobed, granular, greenish to yel- lowish. Digests agar; cellulose not attacked. Fish broth : Flocculent pellicle, turbid, no sediment, fluorescent green. Potato: Abundant, filiform, raised, glistening, smooth, buff, butyrous growth. Nitrites not produced from nitrates. No HiS or indole formed. Plain milk acidified, greenish surface growth. Acid from sucrose. No acid from xy- lose, arabinose, glucose, lactose, manni- tol or rhamnose. Starch is hydrolyzed. Aerobic. Optimum temperature 20° to 28°C; thermosensitive. Source: Isolated from Iridaea curdata. Habitat : On marine algae. Note: Seven additional species are described with as much detail by Angst {loc. cit.) as are the six above species; but he refers to them only as Agarbac- terium Nos. 5, 6, 7, 8, 9, 13, 14, and 15. All digest agar. 8. Flavobacterium uliginosum ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 2G3.) From Latin uligo, ooze or moist mud. Rods: 0.4 to 0.6 by 1.2 to 3.9 microns, some slightly curved, occurring mostlj^ singly with some short chains. Non- motile. Gram-negative. All differential media except the fresh- water broth, litmus milk, and potato were prepared with sea water. Gelatin colonies: 1 mm, orange, sunken. Gelatin stab: Infundibuliform lique- faction. Yellow pigment. Gelatin dis- colored brown. Agar colonies : Sunken, uneven, irregu- lar, gummy colonies which liciuefy agar. Produces orange to yellow pigment and discolors agar brown. Agar slant: Luxuriant, yellowish- FAMILY BACTERIACEAE 631 orange, glistening, filiform, adherent growth which slowh' liquefies agar. Sea-water broth : Dense yellow pellicle, moderate turbidity, slightly viscid sedi- ment. Fresh-water broth: Xo visible growth. Litmus milk: Completelj' decolorized, neutral. Potato: Xo visible growth. Indole not formed. X'itrates rapidly reduced to nitrites. Produces acid but no gas from xylose, glucose, maltose, lactose, sucrose and salicin. Does not ferment glycerol or mannitol. Starch not hydrolyzed. Hydrogen sulfide not formed. Ammonia produced from peptone but not from urea. Casein digested. Fats not hydrolyzed. Agar liquefied rapidly. However, after prolonged laboratory cultivation this organism gradually loses its ability to digest agar. Aerobic, obligate. Optimum temperature 20° to 25°C. Source: jNIarine bottom deposits. 9. Flavobacterium amocontactum Zo- Bell and Allen. (Jour. Bact., 29, 1935, 246.) From Latin amo, to love and con- tactus, touching, contacting. Slender rods: 0.4 to 0.7 by L6 to 2.3 microns, with rounded ends, occurring singly and in irregular clumps. Stain very lightly. Possess well-defined cap- sules. Actively motile by means of peri- trichous flagella. Gram-negative. Gelatin stab: Good filiform growth with rapid saccate liquefaction. Agar colonies: Circular. 2.0 to 4.0 mm in diameter, yellow. Agar slant: Abundant, filiform, smooth, glistening, abundant, bright yellow growth having a but3'rous con- sistency. Originally liquefied agar, but this propertj^ was lost following artificial cultivation. Sea water broth: Good growth with ring at surface. Strong turbidity and abundant viscid sediment. Xo odor. ^lilk: Xo growth. Potato: Xo growth. Potato dialyzed in sea water: Slight yellow growth. Lidole not formed. Xltrites produced from nitrates. Ammonia liberated from peptone. Hydrogen sulfide produced. Xo acid from glucose, lactose, sucrose, xylose or mannitol. Starch not attacked. Optimum reaction pH 8.0. Optimum temperature 18° to 21°C. Facultative aerobe. Distinctive character: Adheres firmly to submerged glass slides; cannot be removed with running water. Source: Many cultures isolated from glass slides submerged in sea water. Habitat: Sea water. Appendix II: Another species de- scribed recently is: Bacillus exedens Wieringa. (Wier- inga, Jour. Microbiol, and Serol., 7, 1941, 121; Bacillus agar-exedens Wier- inga, idem.) From stable manure, leaf- mold and soil. Liquefies agar. 37. Bacterium chitinophilum Hock. (Jour. [Marine Res., 4, 1941, 103.) From ^I. L., chitin and Greek philos, loving. Short rods: 0.35 to 0.65 by 0.95 to 1.5 microns. Motile. Gram-negative. Sea water gelatin: Liquefaction; growth absent in stab but abundant in liquefied zone. Sea water agar plate: Colonies circu- lar, smooth, entire, raised, white. Sea water liquid medium: Moderate growth, sometimes with formation of ring or pellicle. Scant granular sedi- ment. Decomposes natural chitinous material such as horseshoe crab shells and also purified chitin. Four out of five strains produce ni- trites from nitrates. Acid from glucose and usually from sucrose, glycerol and mannitol. One of five cultures produced acid from lactose. Does not digest cellulose. 632 MANUAL OF DETKRMINATIVE BACTERIOLOGY Does not hydrolyze starch. Does not produce hydrogen sulfide. Aerobic. Optimum temperature 20°C. Source: From the shell of a decom- posing horseshoe crab, Limulus polij- phemus, and from the intestinal tracts of Venus mercenaria, Ovalipes ocellatus, Mustelus mustelus and Spheroides ma- culatus. Habitat: Common in marine sand, mud and water. 38. Bacterium chitinochroma Hock. (Jour. Marine Res., 4, 1941, 105.) Short rods: 0.45 to 0.75 by 0.90 to 1.4 microns. Motile. Gram-negative. Sea water gelatin : Active liquefaction ; no growth in stab, but thick bright yel- low growth throughout the liquefied zone. Basic agar plate: Colonies circular, smooth, entire, raised, varying in color from lemon to deep orange. Basic liquid medium: Abundant growth with production of pellicle. Scant granular sediment, increasing with age of culture. Decomposes natural chitinous material such as horseshoe crab shells and also purified chitin. Does not produce nitrites from ni- trates. Acid from glucose and sucrose, but not lactose, glycerol and mannitol. Does not digest cellulose. Hydrolyzes starch. Does not produce hydrogen sulfide. Aerobic. Optimum temperature 20°C. Source: From the intestinal tract of the squid, Loligo pealeii. Common. Habitat: Marine sand, mud and water. Appendix I : The first species of chitin- ovorous bacteria that was described and named was placed in the genus Bacillus because it was a motile rod. 1. Bacillus chitinovorus Benecke. (Bot. Zeitung, 63, 1905, 227.) From M. L. chitin, chitin; varus, devouring. Rods : 0.75 by 2.0 microns. Sometimes in pairs and chains. Motile with peri- trichous flagella. Gram-negative. Gelatin stab: Liquefaction. Mineral agar containing chitin: Good growth if no sugar is added to produce acid. Non-chromogenic. Peptone mineral agar containing chitin : Good growth if reaction is neutral to slightly alkaline. Salt in concentrations up to I5 per cent is favorable for growth. Maximum 4 per cent. Peptone broth: Turbid with heavy, slimy, whitish to brownish pellicle. Nitrites produced from nitrates. Ammonia produced in peptone-chitin media. Acid from glucose and sucrose. Optimum temperature 20°C. Source: Isolated at Kiel from media containing decomposing crab shells and from media containing purified chitin; also from soil. Hal)itat: Brackish water and soil. Notes: Bacillus tumescens Zopf, Ba- cillus cohaerens Gottheil, Bacillus pro- icus vulgaris Kruse, Bacillus coli com- munis Sternberg, Bacillus fluorescens liquefaciens Fliigge, Bacillus megathe- rium De Bary, Vibrio aquatilis Gunther and Spirillum rubrum von Esmarch did not attack chitin under the conditions tested by Benecke (loc. cit.). Benton (Jour. Bact., 29, 1935, 449) describes but does not name 17 types of chitinovorous bacteria isolated from water, mud and plankton of fresh water lakes, from decaying May fly nymph shells, intestinal contents of fish, frogs, bats, snipe, and crayfish. Also shore soil, composts, etc. Twelve types are reported to be monotrichous, two are peritrichous and three, position of flagella not stated. Of two Gram-posi- tive types, one may have been a spore- former and the other a Corynebacterium. Two types digested cellulose. ZoBell and Rittenberg (Jour. Bact., 35, 1938, 275) isolated and studied but FAMILY BACTERIACEAE 633 did not name 31 cultures of chitinoclastic bacteria from marine sources. Out of 16 cultures studied intensively, all were Gram-negative. All but 4 of the 31 cultures were motile. One culture was a coccus and two sjjecies were vibrios. None digested cellulose. *39. Bacterium phosphoreum (Cohn) Molisch. (Micrococcus phosphoreus Cohn, see letter addressed to J. Penn, Verzameling van stukken betreffende het geneeskundig staatstoezicht in Neder- land, 1878, 126; Bacterium phosphores- cens Fischer, Cent. f. Bakt., 3, 1888, 107; Photobacterium phosphorescens Beijer- inck, Arch. N^erl. d. Sci. Exactes, 23, 1889, 401; Streptococcus pJwsphoreus Trevisan, I generi e le specie delle Batteriacee, 1889, 31; Bacillus phos- phoreus Mace, Traits de Bact., Paris, 4th ed., 1901, 995; Micrococcus phos- phorescens Chester, Man. Determ. Bact., 1901, 181; Molisch, Die Leuchtende Pflanzen, 1912, 66; Photobacier phos- phoreum Beijerinck, Folia Microbio- logica. Delft, 4, 1916, 15; Photobacterium phosphoreum Ford, Textb. of Bact., 1927, 615.) Description from Fischer iloc. cit.). Coccobacilli : Occur frequently as zooglea. Xon -motile. Stain lightly with aniline dyes. Gelatin: No liquefaction. Gelatin streak: Gray-white growth. Broth: No growth. Milk: No growth. Potato: No growth. Ferments carbohydrates. Blue-green phosphorescence. Minimum temperature 5°C. Maxi- mum 25°. Optimum for luminescence 10°C. Aerobic, facultative. Source: Isolated from luminous fish. Habitat: Found commonly on dead fish, meat, etc. 40. Bacterium phosphorescens indige- nus (Eisenberg) Chester. (Einheim- ischer Leuchtbaoillus, Fischer, Cent. f. Bakt.; 3, 1888, 107; Photobacterium fischeri Beijerinck, Arch. N6erl. d. Sci. Exactes, 23, 1889, 401; Bacillus fischeri Trevisan, I generi e le specie delle Batteriacee, 1889, 18; Bacillus phos- phorescens indigeniis Eisenberg, Bakt. Diag., 3 Aufl., 1891, 124; Vibrio fischeri Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 342; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 121; Microspira fischeri Chester, Man. De- term. Bact., 1901, 333; Spirillum phos- phorescens Holland, Jour. Bact., 5, 1920, 225; Vibrio phosphorescens Holland, ibid., 226; Achromobacter fischeri Bergey et al.. Manual, 3rd ed., 1930, 220.) Description from Fischer {loc. cit.). Short thick rods: 0.4 to 0.7 by 1.3 to 2.1 microns, with rounded ends, occur- ring singly and in pairs. Motile. Stain with the usual aniline dyes. Johnson, Zworykin and Warren (Jour. Bact., 46, 1943, 167) made pictures with the electron microscope of a culture which they identify with this species. The organisms have a tuft of polar fla- gella, indicating that this species belongs in the genus Pseudomonas. Gelatin stab: Liquefaction. Gelatin colonies: Liquefaction. After one week, circular, 1 mm in diameter, entire. Broth: No growth. ^Nlilk: No growth. Blood serum: No growth. Potato: No growth. Cooked fish: Abundant growth. En- tire surface covered with a gray-white, slimy, phosphorescent mass. Temperature relations: Minimum 5° to 10°C. Optimum 22°C. Aerobic. Source: From sea water at Kiel and from herring. * Dr. Frank H. Johnson, Dept. Bacteriology, Princeton Univ., Princeton, New Jersey, assisted in preparing the section on phosphorescent bacteria, May, 1946 634 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : Live on dead fish and in sea water. 41. Bacterium hemophosphoreum Pfeiffer and Stammer. (Pfeiffer and Stammer, Ztschr. f. Morph. u. Okol. d. Tiere, 20, 1930, 136; Brucella {?) haenw- -phosphor eum Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1033, 67.) Rods: 1.0 by 4.5 microns, the size varying with the medium. Seem to show bipolar staining. Fish agar with 3 per cent sea salt: Good growth. Litmus milk : Acid . Reduction . Potato: Yellow growth, medium be- coming orange. Indole not formed. Nitrites not produced from nitrates. Acid from glucose, sucrose, lactose, maltose, galactose, mannitol and fruc- tose. Phosphorescent. Pathogenic for other insects. Source: Isolated from the blood of diseased larvae of the mealworm, Tene- brio molitor. Habitat: From diseased insect larvae. Appendix 1: The following phospho- rescent species have been described in the literature. Many are incompletely described and they have been placed in various genera without adequate study. Achromobacter argenieophosphorescens (Katz) Bergey ct al. {Bacillus argenieo- phosphorescens I, Katz, Cent. f. Bakt., 9, 1891, 157; Bacterium argenteo-phos- phorescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 121 ; Bacillus argenteo-phosphorescens Migula, Syst. d. Bakt., 2, 1900, 869; Photobacillus I, Miquel and Cambier, Traits de Bact., Paris, 1902, 881; Bergey et al., Manual, 3rd ed., 1930, 221.) From sea water in Elizabeth Bay, Sydney, .Vustralia. Sil- ver-white luminescence. Probably a variety of Photobacterium fischeri Beijer- inck, according to Katz. Achromobacter cyaneophosphorescens (Katz) Bergey et al. {Bacillus cyaneo- phosphorescens Katz, Cent. f. Bakt., 9, 1891, 158; Photobacterium cyaneum Lud- wig, according to Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 331; Photobacterium cyano-phosphorescens (sic) Ford, Textb. of Bact., 1927, 619; Vibrio cyaneo -phosphor escens Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 543; Bergey et al., Manual, 3rd ed. 1930, 221.) From sea water in Little Bay, near Sydney, Australia. Bluish- green luminescence. Identical with or similar to Photobacterium indicum Beijer- inck, according to Katz. Achromobacter luminosum Bergey et al. {Bacillus argenteo-phorphor escens II, Katz, Cent. f. Bakt., 9, 1891, 157; Bac- terium argenteo-phosphorescens Migula, Syst. d. Bakt.. 2, 1900, 435; Photobacil- lus II, Miquel and Cambier, Traits de Bact., Paris, 1902, 882; Bergey et al., Manual, 3rd ed., 1930, 226.) From fish obtained in the market. Greenish -silver luminescence. Achromobacter phosphoreum (Migula) Bergey et al. {Bacillus argenteo-phos- phorescens liqucfaciens Katz, Cent. f. Bakt., 9, 1891, 157; Bacillus phosphoreus Migula, Syst. d. Bakt., 2, 1900, 867; Bergey et al., Manual, 3rd ed., 1930, 222.) From sea water along the coast near Sydnej^ Australia. Luminescence slight. Probably identical with Photo- bacterium luminosum B eijerinckl Achromobacter phosphoricum (Migula) Bergey et al. {Bacillus argenteo-phos- phorescens III, Katz, Cent. f. Bakt., 9, 1891, 157; Bacillus phosphoricus Migula, Syst. d. Bakt., 2, 1900, 870; Photobacillus III, Miquel and Cambier, Traits de Bact., Paris, 1902, 882; Bergey et al., Manual, 3rd ed., 1930, 223.) From cuttlefish {Sepia sp.) obtained in the fish market. Bluish -greenish -white luminescence. ,1 chromobactcr smaragdinophosphores- cens (Katz) Bergey et al. {Bacillus smaragdino-phorphorescens Katz, Cent, f. Bakt., 9, 1891, 159; Bacterium smarag- dino phosphorescens Chester, Ann. Rept. FAIVULY BACTERIACEAE 635 Del. Col. Agr. Exp. Sta., 9, 1897, 124; Bacterium smaragdino-phosphorescens Migula, Sj'st. d. Bakt., 2, 1900, 435; Bac- terium smargadinum (sic) Chester, Man. Determ. Bact., 1901, 181; Bergey et al., ]\Ianual, 3rd ed., 1930, 225.) From her- ring in a fish market in Sydney, Aus- tralia. Green luminescence. Probably identical with Photobacterium phosphor- escens Beijerinck. Bacillus fischeri Dyar. (Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 370; Bacterium fischeri Chester, IMan. Determ. Bact., 1901, 165.) Dyar added to the con- fusion in the nomenclature of phos- phorescent organisms by giving this name to four cultures received by him from the Krai collection labeled Photo- bacterium phosphor escens, Photobac- terium balticum, Photobacterium fischeri and Photobacterium pflilgeri. Bacterium chironomi Issatschenko. (Bulletin du Jardin Imperial botanique a St. Petersbourg, //, 1911, 37; Photo- bacterium chironomi Issatschenko, ibid., 43.) A phosphorescent bacterium from a genus of midges, Chironomus. Bacterium giardi (Kruse) Billet. (Giard and Billet, Compt. rend. Soc. Biol., Paris, 1889, 593; Photobacterium pathogenicum Giard, quoted from Eijk- mann, see abst. in Cent. f. Bakt., 12, 1892, 656; Photobacterium giardi Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 333; Bacillus phosphorescens giardi Kruse, idem; Bacterium phos- phorescens giardi Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 125; Billet, Bull. Sci. France et Belgique, 21, 1898, 144; Bacterium phosphoresccns- giardi Chester, Man. Determ. Bact.. 1901, 182.) Pathogenic for marine crus- taceans. Bacterium hippanici Issatschenko {loc. cit., 47). From fresh water fish. Bacterium luccns (van Tieghem) Niiesch. {Micrococcus lucens van Tieg- hem; Xiiesch, Karsten's Deutsche Flora, 1880; quoted from Ludwig, Cent. f. Bakt., 2, 1887, 375.) From luminous meat. Considered identical with Bac- terium phosphor eum. Bacterium luminosus (Beijerinck) Chester. (Photobacterium luminosum Beijerinck, Arch. N^erl. d. Sci. E.xactes, 23, 1889, 401; Vibrio luminosus Beijer- inck, Bot. Zeit., 1889, 763, according to Trevisan, I generi e le specie delle Batteriacee, 1889, 23; Bacillus luminosus DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 982; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 121; Microspira luminosa Migula, Syst. d. Bakt., 2, 1900, 1015; Photobacter luminosum Beijerinck, Folia Microbio- logica. Delft, 4, 1916, 15.) From sea water. Bacterium pelagia Dubois. (Dubois, Compt. rend. Acad. Sci., Paris, 107, 1888, 502 and 111, 1890, 363; Bacillus pelagia DeToni and Trevisan, in Sac- cardo, Sylloge Fungorum, 5, 1889, 959.) Isolated from the surface of Pelagiae Jioctilucae. Bacterium pfiuegeri Ludwig. (Lud- wig, Ztschr. f. wissensch. Mikrosk., 1, 1884, 181; Micrococcus pflilgeri Ludwig, Hedwigia, Xo. 3, 1884; Arthrobacterium pfliigeri DeBary, 1887; Photobacterium pfiftgeri Beijerinck, Cent. f. Bakt., 8, 1890, 617; Bacterium phosphorescens pflugeri Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 125.) From fish and meat. Considered identical with Bacterium phosphoreum. Bacterium pholas Dubois. (Compt. rend. Acad. Sci., Paris, 107, 1888, 502.) Isolated from Pholadis dactyli. Bacterium phosphorescens Hermes. (Hermes, Sitzungsber. naturf. Freunde, April 19, 1887, quoted from Cent. f. Bakt., 2, 1887, 404; Bacillus hermesi Trevisan, I generi e le specie delle Bat- teriacee, 1889, 18.) From sea water. ]\Iac^ (Traite de Bact., Paris, 4th ed., 1901, 994) says this may be the same as Micrococcus phosphoreus Cohn. Emer- ald-green luminescence. Bacterium phosphorescens gelidus (Eisenberg) Chester. (Phosphoresciren- den Mikroorganismen, Forster, Cent. f. 636 MANUAL OF DETERMINATIVE BACTERIOLOGY Bakt., 2, 1887, 337; Bacillus phosphores- cens gelidus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 182; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 125.) From phosphorescent sea fish. Fischer (Cent. f. Bakt., 4, 1888, 89) states that this organism is the same as his Bac- terium phosphorescens . Coccobacillus acropoma Yasaki and Haneda. (Yasaki and Haneda, 1936; quoted from Harvey, Living Light, Princeton, 1940, 33.) From a fish {Acro- poma japonicum) . Coccobacillus coelorhynchus. (Studied by Hsu, Sei-i-kai Med. Jour., 56, 1937, 1; quoted from Harvej', Annual Rev. of Biochem., iO, 1941, 543.) From a deep- sea fish {Coelorhynchus sp.). Coccobacillus ikiensis. (Quoted from Harvey, Living Light, Princeton, 1940, 263.) Coccobacillus loligo Kishitani. (Kish- itani, Proc. Imp. Acad. Tokj^o, 4, 1928, 69; quoted from Harvey, Living Light, Princeton, 1940, 35.) From the sciuid {Loligo edulis). Micrococcus cijanophos. (Studied by Claren, Ann. d. Chemie, 535, 1938, 122; quoted from Harvej^, Living Light, Princeton, 1940, 184.) Micrococcus physiculus. (Quoted from Harvey, Living Light, 1940, 34.) The cause of luminescence of a fish {Physiculus japonicus) . Microspira phosphoreum. Yasaki. (Yasaki, see Sei-i-kai-zasshi, 45, 1926; quoted from Harvey, Living Light, 1940, 239.) Caused luminescence of a fresh- water shrimp in Japan. Photobacter hollandiae Beijerinck. (Proc. Sect. Sci., Kon. Akad. v. We- tensch., Amsterdam, 3, 1900, 352.) Simi- lar to Pholobacterium luminosum. Photobacter hollandicum Beijerinck. (Folia Microbiologica, Delft, 4, 1916, 15.) Photobacter hollandicum parvum Beijerinck. (Folia Microbiologica, Delft, 4, 1916, 15.) Photobacter splendidum Beijerinck. (Beijerinck, Proc. Sect. Sci., Kon..Akad. V. Wetensch., Amsterdam, 3, 1900, 352; Vibrio splendidus Lehmann and Neu- mann, Bakt. Diag., 7 Aufl., 2, 1927, 543; Photobactcrium splendidum, quoted from Harvey, Living Light, Princeton, 1940, 204.) Maj' be a variety of Photobac- terium indicum. Photobacter splendor maris Beijerinck. (Proc. Sect. Sci., Kon. Akad. v. We- tensch., Amsterdam, 3, 1900, 352.) May be a variety of Photobacterium indicum. Photobacterium Beijerinck. (Beijer- inck, Arch. N^erl. d. Sci. Exactes, 23, 1889, 401; Photobacter Beijerinck, Proc. Sect. Sci., Kon. Akad. v. Wetensch., Amsterdam, 3, 1900, 352; Photospirillum Miciuel and Cambier, Traits de Bact., Paris, 1902, 888; Pholomonas Orla-Jen- sen, Jour. Bact., 6, 1921, 271.) Photo- bacterium phosphorescens is the type species of this genus. See Bacterium phosphoreum. Several species are placed in this genus by Fischer. Photobacterium annulare Fischer. (Fischer, Ergebnisse d. Plankton-Ex- pedition d. Humboldt-Stiftung, 4, 1894, 41; Microspira anrmlaris Migula, Syst. d. Bakt., .g, 1900, 1014.) From sea water. Photobacterium balticum Beijerinck. (Einheimischer Leuchtbacillus, Fischer, Cent. f. Bakt., 3, 1888, 105; Beijerinck, Akad. V. Wetenschappen, Afdeel. Na- tuurk., 2de Reeks, 7, 1890, 239; see abst. in Cent. f. Bakt., 8, 1890, 617; Vibrio balticus Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 341.) From water of the Baltic Sea. The relationship of Photobacterium balticum to Bacterium phosphorescens indigenus is not clear. The former species is based on a culture sent by Fischer to Beijerinck labeled Einheimischer Leuchtbacillus which Beijerinck considered to be different from his Photobacterium fischeri. Photobacterium caraibicum Fischer. (Fischer, loc. cit., 1894, 41; Microspira caraibica Migula, loc. cit., 1015.) From sea water. Photobacterium coronatum Fischer. (Fischer, loc. cit., 4:1; Microspira coronata FAMILY BACTERIACEAE 63: Migula, loc. cit., 1013.) From sea water. Photohacterium degenerans Fischer. (Fischer, loc. cit., 37; Microspira de- generans Migula, loc. cit., 1015; Bacillus degenerans Beijerinck, Folia Microbio- logica, Delft, I, 1912, 1.) From sea water. Photobacterium delgadense Fischer. (Fischer, loc. cit., 37; Microspira delga- densis Migula, loc. cit., 1014.) From sea water. Photohaclerium glutinosum Fischer. (Fischer, loc. cit., 41; Microspira gluli- nosa Migula, loc. cit., 1014.) From sea water. Photobacterium hirsutum Fischer (loc. cit., 41). From marine fish. Photobacterium papillare Fischer. (Fischer, loc. cit., 41; Microspira papil- laris Migula, loc. cit., 1016.) From sea water. Photobacterium sepiae. (Quoted from Doudoroff, Jour. Bact., U, 1942, 451, who obtained a culture so labeled which had come from Prof. Kluyver's collec- tion at Delft.) Photobacterium tuberosum Fischer. (Fischer, loc. cit., 37; Microspira tubcrosa Migula, loc. cit., 1014; Photobacter tuberculatum Beijerinck, Folia Micro- biologica, Delft, 4, 1916, 15.) From sea water. Pseudomonas toyameusis. (Quoted from Harvey, Living Light, Princeton, 1940, 263.) Sarcina noctiluca Heller. (Heller, Arch. f. Physiol., path. Chem. u. Mikr., N.F., 6, 1853-54, 44; see Harvey, Living Light, Princeton, 1940, 6.) From fish. Possibly the same as Bacterium phos- phoreum Molisch. 42. Bacterium erythrogloeum Ruhland and Grohmann. (Cent. f. Bakt., II Abt., 61, 1924, 256.) From Greek erij- thros, red and glota, glue. Rods : 0.5 b}- 2.0 microns. Non-motile. Gram-negative. Gelatin plate: Red, droplet-like colonies. Gelatin stab: No liquefaction. Agar plate: Red, droplet-like colonies. Agar slant: Raised, non-spreading, glistening, brick-red growth. Potato: Abundant, brick-red, warty. Aerobic. Facultative autotroph. O.xidizes hydrogen in an inorganic medium under an atmosphere of Ho, O2, and CO2. Produces a pellicle on the inorganic liciuid medium. Source: Calcareous soil. Habitat: Probably widely distributed in soil. 43. Bacterium lentulum Grohmann. (Cent. f. Bakt., II Abt., 61, 1924, 256.) Rods : 0.5 by 1 to 2 microns. Motile by long thin peril richous flagella. Gram- negative. Gelatin plate: Colonies appear like milk droplets. Gelatin stab: No liquefaction. Agar plate: Tough, ochre yellow col- onies about 7 mm in diameter. Agar streak: Parchment -like, folded, yellow streak about 1 cm broad. Potato: Heavy, yellow growth. Aerobic. Facultative autotroph. Oxidizes hydrogen in an inorganic medium under an atmosphere of H2, O2, and CO 2. Produces a heavy folded pellicle on the inorganic liquid medium. Source: Soil poor in lime. Habitat: Probablj' widely distributed in soil. 44. Bacterium leucogloeum Ruhland and Grohmann. (Cent. f. Bakt., II \ht., 61, 1924, 256.) From Greek leukos, white and glota, glue. Rods: 0.5 by 0.7 to 52 (?) microns. Motile by means of peritrichous flagella. Gelatin stab: No liquefaction. Agar streak: Wide, slimy, wet, ivory- colored growth. Potato: Gray-brown slime. Aerobic. Facultative autotroph. Oxidizes hydrogen in an inorganic medium under an atmosphere of H2, O2, 638 IVIANUAL OF DETERMINATIVE BACTERIOLOGY and CO 2. Produces a pellicle on the inorganic liquid medium. Source: Calcareous soil. Habitat: Probably widely distributed in soil. *4o. Bacteriiun stewartii Erw. Smith. (Sweet corn bacillus, Stewart, N. Y. Agr. Exp. Sta. Bui. 130, 1897, 423; Pseudomo7ias stewarli Smith, Proc. A. A. A. Sci., 47, 1898, 422; Smith, Pact, in Rel. to Plant Dis., 3, 1914, 89; Aplano- bacter stewarli McCulloch, Phytopath., 8, 1918, 440; Bacillus stewarli Holland, Jour. Pact., 5, 1920, 220; PMjtomonas stewartii Bergey et al., Manual, 1st ed., 1923, 192.) Named for F. C. Stewart, American plant pathologist. Description from Smith, U. S. Dept. Agric, Div. Veg. Phys. and Path., Bui. 28, 1901. Rods: 0.4 to 0.7 by 0.9 to 2.0 microns. Capsules. Non-motile (McCulloch, loc. cit.). Gram-negative. Gelatin: No liquefaction. Nutrient agar colonies: Small, round, yellow colonies. Broth: Growth feeble with whitish ring and j'ellow precipitate. Milk: Yellow ring but no visible action on the milk. Slightly acid. Nitrites not produced from nitrates. McNew (Phytopath., 28, 1938, 773) states that less virulent strains assimi- late only organic nitrogen; those of intermediate virulence assimilate nitro- gen from inorganic salts without reduc- tion of nitrates to nitrites; virulent strains reduce nitrates to nitrites. H3-drogen sulfide not formed. Indole production slight or none. Reduction of methylene blue in Dun- ham's solution feeble or doubtful. Acid but no gas from glucose, galac- tose, sucrose, mannitol and glycerol. No acid from maltose. Acid from fruc- tose, arabinose and xylose (McNew, loc. cit.). Starch not hydrolyzed. Optimum temperature 30°C. Maxi- mum 39°C. Minimum 8°C. Optimum pH 6.0 to 8.0. Limits about pH 4.5 to 8.5. 8 per cent salt restricts growth. Strict aerobe. Source: From wilted sweet corn. Habitat: Pathogenic on corn, Zea mays. Sweet corn very susceptible and field corn slightly so. 46. Bacterium tardicrescens Mc- Culloch. (McCulloch, Phytopath., 27, 1937, 135; Phytomonas tardicrescens Burkholder, Phytopath., 27, 1937, 617.) From Latin, slow growing. Rods: 0.6 to 0.8 by 1.58 microns. Mo- tile with a polar flagellum. Gram-nega- tive. Gelatin: No liquefaction. Beef-extract agar colonies: Circular, mustard yellow, edges entire, 1 to 1.5 mm in diameter. Broth: Light clouding. Milk: Slightly alkaline. Clearing after 5 to 6 weeks. Nitrites are produced from nitrates. Indole not produced. No HoS produced or feebly so. Acid but no gas from glucose, fructose, galactose, arabinose, xylose and rham- nose. Alkaline reaction from salts of citric, malic and succinic acid. Starch is not hydrolyzed. Not lipolytic (Starr and Burkholder, Phytopath., 32, 1942, 603). Optimum temperature 26°C. Maxi- mum 32°C. Minimum 5°C (McCulloch, Phytopath., 28, 1938, 648). Optimum pH 6.5 to 7.5. Growth slight at 5.8 and 8.0 (McCulloch, loc. cit.). No growth with 3 per cent salt (Mc- Culloch, loc. cit.). Aerobic. Distinctive character: Very slow grower. *The section covering species of interest to plant pathologists has been pre- pared by Prof. Walter H. Burkholder, Cornell Univ., Ithaca, New York, May, 1946. FAMILY BACTERIACEAE 639 Source : Isolated by McCulloch and by Burkholder from blighted iris leaves. Habitat: Pathogenic on Iris spp. 47. Bacterium albilineans Ashby. (Ashby, Trop. Agr., Trinidad, 6, 1929, 135; Phytomonas albilineans JMagrou, in Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 326.) From Latin, produc- ing white streaks. Description taken from Martin, Car- penter and Weller, The Hawaiian Planters' Record, 36, 1932, 184. . Rods: 0.25 to 0.3 by 0.6 to 1.0 micron, occurring singly or in chains. Motile with a polar flagellum. Gram-negative. Agar colonies: After 7 to 10 days, minute transparent drops, moist, shin- ing. Honey yellow to Naples yellow. Gelatin: No liquefaction. Milk: Growth, but no visible change in the milk. No growth with ammonium salts, ni- trates, or asparagine as a source of ni- trogen. No growth in peptone water without carbohydrates. Invertase secreted. Starch is not hydrolyzed. Optimum temperature about 25°C. Maximum 37 °C. Distinctive characters: Differs from Xanthomonas vascularum which produces a large gummy type of colony, and which is a very active organism biochemically. The two pathogens also differ in the type of lesion they produce on .sugar cane. Source: Isolated by D. S. North (Co- lonial Sugar Ref. Co., Sidney, N.S. Wales, Agr. Rept., 8, 1926, 1) from white stripe and leaf scald of sugar cane in Australia. Habitat : Vascular pathogen of sugar cane, Saccharutn officinarum. Appendix I: The following species liave been described from diseased plant tissues but may not, in some cases at least, have been the cause of the disease. Bacillus betae Migula. (Kramer, Oes- terreich. landwirtsch. Centralb., 1891, Heft 2 and 3; Migula, Syst. d. Bakt., 2, 1900, 779.) The cause of a disease of the sugar beet (Beta vulgaris). Bac. caryophyllacearum Dufrenoy. (Compt. rend. Soc. Biol., Paris, 81, 1918, 920; probably there is an earlier reference to this species.) On Dianthus, Sapo- nnria and Lychnis. Bacillus coffeicola Steyaert. (Rev. Zoo. et Bot. Afr., 22, 1932, 137.) From nodules on coffee roots. Bacillus lacerans Migula. {Bacillus a, Busse, Ztschr. f. Pflanzenkr., 7, 18 — , . 72; Migula, Syst. d. Bakt., 2, 1900, 780.) From diseased sugar beets. Bacillus maculicola Delacroix. (Dela- croix, Compt. rend. Acad. Sci. Paris, HO, 1905, 680; Bacterium maculicola Stapp, in Sorauer, Handb. d. Pflanzen- krankheiten, 5 Aufl., 2, 1928, 276; Aplan- obacter maculicola Elliott, Manual Bact. Plant Path., 1930, 8; Phytomonas nico- tianae-tabaci MagTou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 386.) From diseased spots on leaves of tobacco. Bac. nucleophyllus Dufrenoy. (Compt. rend. Soc. Biol., Paris, 81, 1918, 920, nomen nudum.) On Rhododendron fer- rugineum. Bac. trilici Dufrenoy. (Compt. rend. Soc. Biol., Paris, 81, 1918, 920, nomen nudem; not Pseudomonas tritici Hutchin- son, India Dept. of Agr., Bact. Ser., 1, 1917, 174.) On wheat. Bacillus vitis Montemartini. (Rev. Patol. Veg., 6, 1913, 175.) Pathogenic on the grape (Vitis vinifera). Bacterium apii Brizi. (Lav. e Relaz. d. Reg. Staz. di Patol. Veg., Roma, Gennio-Giugno, 15, 1896 and Atti R! Ac- cad. Naz. Lincei, Rend. CI. Sc. Fis., Math, e Nat., Ser. 5, 6, 1897, 233.) Motile. From rot of celery. Bacterium betae Chester. (Bacterial parasite, Arthur and Golden, Indiana Agr. Exp. Sta., Bull. 39, 1892, 61; Ches- ter, Ann. Rept. Del. Col. Agr. E.\p. Sta., 9, 1897, 128; Bacillus arthuri Migula, Syst. d. Bakt., 2, 1900, 681.) Motile. From diseased sugar beet tubers. Bacterium briosianum Pavarino. 640 MANUAL OF DETERMINATIVE BACTERIOLOGY (Atti del. R. Accad. Naz. Lincei, Rend. CI. Sci. Fis., Math, ot Nat., W, 1911, 161.) Motile. From lesions on the vanilla vine. Bacterium casiiniculum Cavara. (Rev. d. Pat. Veg., 7, 1914, 5.) Motile. From chestnut canker. Bacterium corylii Brzezinski. (Bull. Intern. Acad, des Sci. Cracovie, CI. Sci. Math, e Nat., 1903, 139.) Motile. From diseased filbert trees. Bacterium dendrobii Pavarino. (Rev. di Pat. Veg., 5, 1912, 242.) Bacterium dianthi Chester. (Para- sitic bacteria, Arthur and Bolley , Purdue Univ. Agr. E.xp. Sta., Bull. 59, 1896, 21; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 106; Bacillus dianthi Chester, Man. Determ. Bact., 1901, 253; Pseudomonas dianthi E. F. Smith, U. S. Dept. Agr., Div. Veg. Phys. and Path., Bull. 28, 1901, 153.) Motile. From lesions on carnation leaves. Bacterixmi fici Cavara. .(1st. Bot. del. R. Univ. di Catania, Atti Acad. Gioen., 18, Mem. 14, 1905, 1; Phrjtomonas (?) fici Magrou, in Hauduroy et al., Diet. d. Bact. Path., 1937, 354.) Motile. Causes a blight of figs. Bacterium lycopersici var. vitiati Strzalkowska. (Strzalkowska, Acta Soc. Bot. Poloniae, Warsaw, 7, 1930, 611; Phijtomonas vitiati Burkholder, in Man- ual, 5th ed., 1939, 216.) From rotting tomato. Bacterium mali Brzezinski. (Bull. Intern. Acad. Sci. Cracovie, CI. Sci. Math, e Nat., 1903, 100.) Motile. From apple canker. Bacterium montemartinii Pavarino. (Rev. di Pat. Veg., 5, 1911, 65.) Motile. From wisteria canker. Bacterium {?) oncidii Peglion. (Peg- lion, 1899, quoted from Hauduroy et al.. Diet. Bact. Path., 1937, 388; Bacillus oncidii Stevens, 1913; Phytomonas {?) oncidii Hauduroy et al., idem.) From an orchid (Oncidium sp.). Bacterium pini Chester. (Bacillus des tumeurs du Pin d'Alep, Vuillemin, Compt. rend. Acad. Sci., Paris, 107, 1888, 874 and 1184; Bacillus vuillemini Trevi- san, I generi e le specie delle Batteriacee, 1889, 19; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 127; Pseudo- monas pini Petri, Ann. 1st. Supt. For. Naz. Firenze, 9, 1924, 187.) From galls on pine (Pinus halepensis) . Bacterium putredinis Davaine. (Da- vaine, Bact^ries, in Dictionnaire En- cyclop^dique des Sci. M^dicales, 1866; Bacillus putredinis Trevisan, Add. ad Gen., p. 36; see DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1025; "not Bacillus putredinis Weinberg, Nativelle and Pr^vot, Les Microbes Ana^robies, Paris, 1937, 755.) Causes a soft rot of several plants. Bacterium pyri Brzezinski. (Bull. Internat. d. I'Acad. des Sci. de Cracovie, CI. Sci. Math, e Nat., 1903, 130.) Mo- tile. From pear canker. Bacterium rubefaciens Burr. (Burr, Ann. App. Biol., 15, 1928, 570; Phyto- monas rubefaciens Magrou, in Hauduroy et al.. Diet. Bact. Path., 1937, 406; not Bacterium rubefaciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 115.) Bacterium suberfaciens Burr. (Burr, Ann. App. Biol., 15, 1928, 570; Phijto- monas suberfaciens Magrou, in Hauduroy et al., Diet. Bact. Path., 1937, 417.) Motile. From diseased potato tubers. 48. Bacterium rubefaciens (Zimmer- mann) Chester. {Bacillus rubefaciens Zimmermann, Die Bakterien unserer Trink- und Nutzwasser, Chemnitz, /, 1890, 26; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 115; Erythro- baciUus rubefaciens Holland, Jour. Bact., 5, 1920, 223; Serraiia rubefaciens Bergey et al., Manual, 1st ed., 1923, 92; Chromo- bacterium rubefaciens Topley and Wilson, Princ. Bact. and Immun., 1, 1931, 402.) From Latin ruber, red and /act'o, to make. Rods: 1.0 to 1.6 microns in length, oc- curring singly and in pairs. Actively motile. Gram-negative. Gelatin colonies: Minute, white. Gelatin stab: Surface growth yellow- ish, the medium taking on a red tinge. No liquefaction. FAMILY BACTERIACEAE 641 Agar colonies: Small, white, with erose margin. Agar slant: White, smooth, glistening, somewhat luxuriant, the medium taking on a wine red color. Broth: Turbid with white pellicle, the medium slowly assuming a reddish tinge. Litmus milk: Acid, with slow coagula- tion and reduction of the litmus. Be- coming alkaline. Potato: A heavy, white, creamy layer, which later becomes yellowish -brown. Indole not produced. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 25°C. No growth at 37°C. Habitat : Water. 49. Bacterium rubidum (Eisenberg) Chester. {Bacillus rubidus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 88; Bacterium rubidus (sic) Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 107 and 115; Serratia rubida Levine and Soppeland, Iowa State Coll. Engineering Exp. Sta. Bull. 77, 1926, 53.) From Latin rubidus, red. Description fnmi Eisenberg (loc. cil.). Levine and Soppeland (loc. cit.) found an organism in buttermilk which they identified as Serratia rubida. Their de- scription is more complete than that given by Eisenberg but differs from the original in several respects. Rods : Medium size with rounded ends, often in long chains. Motile. Gelatin colonies: Circular, finely gran- ular, entire, with reddish center. Slow growth. Gelatin stab: Liquefaction. Brown- ish-red sediment. Agar colonies: Small, flat, smooth, amorphous, entire, brownish-red. Slow growth . Agar slant: Brownish-red streak. Spreading over surface. Potato: Brownish -red growth. Blood serum liquefied, red pigment. Aerobic, facultative. Does not grow well at 37°C. Source: Water. 50. Bacteritim latericeum (Adametz) Lehmann and Neumann. (Bacillus la- ter iceus Adametz, Die Bakterien der Trink- und Nutzwiisser, Mitteil. der oestrr. Versuchsanst. f. Brauerei u. Malzerei in Wien, 1888, 50; Bacillus erythraeus Trevisan, I generi e le specie delle Batteriacee, 1889, 19; Bacterium latericeum Lehmann and Neumann, Bakt. Diag., 1 Aufi., 2, 1896, 258; Serratia latericea Bergey et al., Manual, 1st ed., 1923, 94.) From Latin latericeus, brick. Rods: 0.5 to 0.7 by 1.0 to 1.3 microns. Non-motile. Gram-negative. Gelatin colonies: Small, white, granu- lar, with slightly irregular margin. Gelatin stab: A thin, dry, spreading, cream -pink surface growth. No lique- faction. Agar colonies: Dry, glistening, whit- ish, with irregular margin. Agar slant: Brick-red, smooth, glis- tening, butyrous. Broth: Thick pellicle; fluid clear. Litmus milk: Alkaline. Potato: Brick-red streak. No gas from carbohydrate media. Indole not produced. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 25° to 30°C. Habitat: Water. 51. Bacterium alginicum Waksman, Carey and Allen. (Jour. Bact., 28, 1934, 213.) Rods short to almost spherical, 0.6 to 1.0 micron in diameter. Sluggishly motile. Capsule-forming. Gram- negative. Alginic acid plate: White, finely granulated colonies, with entire margin. Does not clear up the turbiditj" in plate. Odor formed, resembling that of old potatoes. Alginic acid liquid medium: Thin pellicle, weak alginase formation. Sea water gelatin : Thin growth throughout gelatin stab, no liquefaction in 7 days at 1S°C. Agar liquefaction: None. Sea water glucose broth: Uniform but 642 MANUAL OF DETERMINATIVE BACTERIOLOGY very limited turbidity; no pellicle; no sediment. Litmus milk containing salt: No ap- parent growth. Potato moistened with sea water: Moist, spreading growth, cream-colored; heavy sediment in free liquid at bottom. Starch plate: Limited, pale blue growth; no di9,stase. Aerobic. Optimum temperature 20°C. Source: From sea water, and from the surface of algal growth. Habitat: Common in sea water. 52. Bacterium terrestralginicum Waks- man et al. (Waksman, Carey and Allen, Jour. Bact., 28, 1934, 217.) Long rods, with somewhat rounded ends, usually single, but also in pairs, and occasionally in chains of shorter rods. 1.0 to 1.5 by 1.5 to 2.5 microns. Motile. Granular. Gram-negative. Alginic acid plate: Colonies small, whitish in appearance with a slight metallic sheen. Alginic acid liquid medium: Medium at first clouded. Later, a pellicle is formed on the surface of the medium, which is soon broken up due to active gas formation. Reaction of medium becomes slightly alkaline. Gelatin medium : Slow growth through- out stab, slow liquefaction at surface of medium at 18°C. Agar liquefaction: None. Glucose broth: Abundant turbidity, some sediment, no pellicle, slightly fluorescent. Litmus milk: Acid, milk coagulated, only limited digestion of coagulum. Potato: Abundant, pinkish, compact, dry growth on surface of plug, the rest of plug becoming gray, with a tendency to darkening. Starch plate: Limited growth along streak, no diastase. Aerobic to facultative anaerobic. Optimum temperature 30°C. Source: From New Jersey soil. Habitat: Soil. 53. Bacteriimi cygni Migula. (Septi- kamiebacillus der Schwane, Fiorentini, Cent. f. Bakt., 19, 1896, 935; Migula, Syst. d. Bakt., 2, 1900, 365; Bacillus cygneus Chester, Manual Determ. Bact., 1901, 221.) From Latin cygnus, swan. Rods: Motile. Gram-negative. This organism may have been the fowl cholera or septicemia organism (Pasteurella avicida Trevisan); but. is more probably closely related to the organism which causes keel in ducklings {Salmonella anatis Rettger and Scoville). Source: From a swan. Habitat: The cause of an infectious disease of swans in the city park at Milan, Italy in 1895. 54. Bacterium cyprinicida Plehn. (Plehn, Cent. f. Bakt., I Abt., Orig., 35, 1903-04, 461; Klebsiella cyprinicida Bergey et al., Manual, 2nd ed., 1925, 266.) From Greek kyprmos, carp and Latin caedo, to kill. Rods: 0.8 by 1.0 micron, occurring singly and in chains. Capsulated. Non-motile. Gram-negative. Gelatin colonies: White, glistening, conve.x, with slight fluorescence around the colony in three or four days. Gelatin stab: White, convex surface growth. No liquefaction. Agar slant: White, glistening layer, becoming slimy. Broth: Turbid, with thick gray pellicle and slimy sediment. Litmus milk: Slightly alkaline. No coagulation. Potato: Light yellowish layer, be- coming dark brownish. The medium is dark violet-gray. Indole not formed. Nitrites not produced from nitrates. No acid from carbohydrate media. Aerobic, facultative. Optimum temperature 10° to 20°C. Habitat: The cause of a fatal disease in carp, showing as red spots on the ventral surface. 55. Bacteriiunparvulum Conn. (N.Y. FA^^LY BACTERIACEAE 643 Agr. Exp. Sta. Bui. -194, 1922,26.) From Latin, very small. Very small rods: 0.1 to 0.2 by 0.3 to 0.5 micron. Xon-motile. Gram-nega- tive. Gelatin plate: Punctiform colonies. Agar plate: Punctiform colonies. Grows poorly in liquid media. Indole not formed. Nitrites produced from nitrates. No acid from glucose, lactose, sucrose, glycerol or ethyl alcohol in either liquid or solid media. Starch not digested. Optimum temperature 25°C. Stricth' aerobic. Distinctive character: Causes strong volatilization of ammonia from a mix- ture of horse feces and urine. Source: From manure. Habitat: Soil. 56. Bacteriiun methylicum (Loew) Mig- ula. {Bacillus methylicus Loew, Cent, f. Bakt., 12, 1892, 465; Migula, Syst. d. Bakt., 2, 1900, 447.) From the chemical term, methyl. Short, thick rods: 1.0 by 2.0 to 2.5 microns. Gram stain not recorded. Gelatin colonies: After 2 days, round to oval, yellowish, entire; later edges ciliate. Liquefaction. Glucose gelatin stab: Liquefaction crateriform. Whitish-yellowish sedi- ment. No liquefaction in depth. Glucose gelatin stab: In depth, little or no growth, slowly liquefied near surface. Agar stab: Surface growth spreading, grayish -white. No growth in depth. Broth: No turbidity. On the surface and adherent to the walls, a white ring which precipitates on shaking. Potato: Growth very slow, pure white, adherent. Grows well in 0.5 per cent methyl alcohol, 0.05 per cent di calcium phos- phate, and 0.01 per cent magnesium sul- fate, on which broth it forms a reddish pellicle. Possesses the ability to decompose formaldehyde and formic acid salts with formation of a reddish pellicle. Aerobe. Source: A culture contamination from the air. Habitat: Probably soil. Appendix I: A few of the numerous Gram-negative, motile or non-motile, non- spore-forming rods that do not belong in the groups previously listed in this genus are described here. All have been placed in the genus Bacillus by those who have described them, although none form spores. I. Produce a pink to red chromogenesis. A. Motile. 1. Bacillus lactorubefaciens. B. Non-motile. 1. Gelatin liquefied. 2. Bacillus rubricus. 3. Bacillus rujus. 2. Gelatin not liquefied. a. Salmon pink on agar. 4. Bacillus mycoides corallinus. aa. Vinous red on agar. 5. Bacillus bruntzii. II. Produces a water-soluble orange to emerald green pigment. A. Motile. 1. Gelatin liquefied. 6. Bacillus aurantiacus tingitanus 644 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Bacillus lactorubefaciens Gruber. (Gruber, Cent. f. Bakt., II Abt., 8, 1902, 457; Serralia lactorubefaciens Ber- gey et al., Manual, 1st ed., 1923, 92.) From Latin, to make milk red. Small rods: 0.4 to 0.6 by 3.5 microns, occurring singly and in pairs. Motile with peritriclious flagella. Gram reac- tion not given. Gelatin colonies: Grayish-white, smooth, glistening, spreading. Gelatin stab: At times arborescent; the medium tinged with red. No lique- faction. Agar colonies : Circular, lobed, grayish, contoured. Agar slant: White, spreading growth. Broth: Turbid, with grayish pellicle and slimy sediment. Limus milk: Becomes rose red, slimy, slightly acid, without coagulation. Potato: White, spreading growth. No gas from carbohydrate media. Indole not produced. Nitrites produced from nitrates. Aerobic, facultative. Optimum temperature 25°C. Habitat: Milk. 2. Bacillus rubricus Hefferan. (Hef- feran. Cent. f. Bakt., II Abt., 11, 1903, 403; Erythrohacillus rubricus Holland, Jour. Bact., 5, 1920, 220; Serralia rubrica Bergey et al.. Manual, 1st ed., 1923, 313; Chromobacterium rubricum Topley and Wilson, Princ. Bact. and Immun., 7, 1931, 402.) Rods: 0.7 to 0.9 by 1.0 to 4.0 microns, occurring singly. Non-motile. Gram reaction not» given. Gelatin colonies: Small, circular, yel- low-orange, deepening to red. Gelatin stab: Slow liquefaction. Old cultures lose this property. Agar colonies: Circular, raised, entire. Agar slant: Moist, spreading, white to pink, gradually deepening in color. Broth: Turbid, with viscid sediment. Litmus milk: Alkaline. Potato: Slight growth, blight pink, turning coral red. Indole not produced. Nitrites not produced from nitrates. No acid or gas from carbohydrate media. Aerobic, facultative. Optimum temperature 25° to 30°C. No growth at 37 °C. Source: Isolated from Mississippi river water, also from buttermilk. 3. Bacillus rufus Hefferan. (Hef- feran, Cent. f. Bakt., II Abt., 11, 1903, 313; Erythrobacillus rufus Holland, Jour. Bact., 5, 1920, 220; Serralia rufa Bergey et al., Manual, 1st ed., 1923, 95.) From Latin rufus, red. Differs from Bacillus rubricus in show- ing more luxuriant growth on potato and slower action in milk. Source: From Mississippi Uiver water. 4. Bacillus mycoides corallinus Hef- feran. (Hefferan, Cent. f. Bakt., II Abt., 11, 1903, 459; Serralia corallina Bergey et al., Manual, 1st ed., 1923, 93.) Small, slender rods: 1.2 to 2.0 microns in length, occurring singly and in pairs. Non-motile. Gram reaction not given. Gelatin colonies: Minute, becoming pink, smooth, raised. Gelatin stab: Slow growth. Raised, smooth, glistening, pink surface growth. Fine, feathery growth in stab. No lique- faction. Agar colonies: Minute, with filamen- tous margin. Agar slant: Smooth, moist, salmon pink. Broth; Turbid, with pink flakes on surface. Litmus milk: Alkaline, with red sur- face. Potato: Like agar slant. Indole not formed. Nitrites produced from nitrates. No gas from carbohydrate media. Aerobic, facultative. Optimum temperature 25° to 30°C. Source: Isolated from Mississippi river water. 5. Bacillus bruntzii Nepveux. (Nep- veux, Compt. rend. Soc. Biol., Paris, 72, 1920, 242; These, Fac. de Pharm., Jouve et Cie, Paris, 1920. 136 pp.; Serralia bruntzii Bergey et al., Manual, FAMILY BACTERIACEAE 645 3rd ed., 1930, 125.) Xamcd for Prof. Bruntz of Paris. Bacillus roseus fluorescens Mar- chal (Trav. Lab. Microbiol. Fac. Pharm. Xanc}-, 1937, 90) is regarded by Lasseur (personal communication, 1938) as identical with Bacillus hruntzii Xepveux. Rods: 0.3 to 0.5 by 1.25 to 1.5 microns, occurring singly and in pairs. Xon- motile. Gram-negative. The cells store volutin and glycogen as reserve materials. Gelatin colonies: Circular, gray, smooth, contoured, glistening, undulate margin, becoming red. Gelatin stab: Xo liquefaction. Agar colonies: Circular, flat, smooth, contoured, radiate margin, vinous red. Agar slant: Smooth, echinulate, bu- tyrous, vinous red in color. Broth: Turbid. Litmus milk: Unchanged. Indole not formed. Xitrites produced from nitrates. Acid from glucose, fructose, maltose, lactose, sucrose, mannitol, dulcitol and glycerol. Aerobic, facultative. Optimum temperature 20° to 25°C. Habitat: Water. 6. Bacillus aurantiacus tingitanus Remlinger and Bailly. (Compt. rend. Soc. Biol. Paris, 119, 1935, 246.) Short rods: Usually 2 to 3 microns, sometimes 5 to 6 microns long. Activeh^ motile. Gram-negative. Growth occurs on all the ordinary nutrient media. Fluorescent bright orange pigment. Gelatin: Rapid liquefaction. Milk: Slow coagulation. Synthetic broth: Lasseur, Dupaix- Lasseur and jNIarion (Trav. Lab. Micro- biol. Fac. Pharm. Xancy, Fasc. 9, 1936, 34) recognize two rough types of this organism, one of which forms a smooth and the other a wrinkled pellicle. The smooth type gives a rough (pH 4.7) or a smooth (pH 6.3) pellicle according to the pH of the medium. Indole not formed. Artichoke media: Luxuriant growth. Emerald green pigment produced. On transferring the culture to potato, the bright orange pigment reappears. Coagulated serum: Xo liquefaction. Acid from sucrose, lactose, glucose, mannitol and maltose. Xon -pathogenic. Optimum pH 6.6. Xo growth at pH 6.2, but grows at pH 7.8. Optimum temperature 20°C. Good growth from 15° to 37 °C. Aerobic. Pigment: Orange or capucine pigment which diffuses throughout the medium. Xot affected by the presence or absence of light. Pigment production depends on the growth of the culture, not on the acidity of the medium. Insoluble in acetone, amyl alcohol and gasoline. Partially soluble in ether and ethyl alcohol which are colored yellow. Distinctive character: A fluorescent pigment of an unusual shade (bright orange). Source: From water at Tangiers. Habitat: Presumably water. Appendix II :* The anaerobic genus Methanobacterium was proposed tentatively by Kluyver and Van Xiel in 1936 with indication that they regarded Sohngen's meth- ane bacterium as the type species of the genus. Later, Barker (1936) found or- ganisms that he regarded as identical with those previously isolated by Sohngen and he proposed the name Methanobacterium sohngenii for this species. A second species found at the same time was named Methanobacterium omelianskii and it was identified as the species previously described but not named by Omeliansky. At the time, he felt that these anaerobes should be included in the family Mycobacteriaceae (1936, * The manuscript for this section has been reviewed by Dr. H. Albert Barker, University of California, Berkeley, California, February, 1945. 646 MANUAL OF DETERMINATIVE BACTERIOLOGY p. 422). In 1940, he discovered that the second species produced spores. In a per- sonal communication (March 20, 1945) he suggests that further work is needed before the relationships of these organisms can be clarified. Genus A. Methanobacterium Kluyver and Van Niel. (Cent. f. Bakt., II Abt., 94, 1936, 399.) Straight or slightly bent rods, sometimes united in bundles or long chains. Usu- ally non-motile. Endospores sometimes formed. Anaerobic. Chemo-heterotro- phic or chemo-autotrophic oxidizing various organic or inorganic compounds and reducing carbon dioxide to methane. Gram- variable, usually negative. The type species is Methanobacterium soehngenii Barker. 1. Methanobacteriimi soehngenii Bar- ker. (Methane bacterium, Sohngen, Dis- sertation, Delft, 1906; Barker, Arch. f. Mikrobiol., 7, 1936, 433.) Named for Prof. X. L. Sohngen who first studied this organism. Rods : Straight to slightly bent, moder- ately long. Non-motile. Non-spore- forming. Gram-negative. In liquid cultures cells are character- istically joined into long chains which often lie parallel to one another so as to form bundles. Acetate and n-butyrate but not pro- pionate are fermented with the produc- tion of methane and carbon dioxide. Ethjd and n-butyl alcohols not fer- mented . Obligate anaerobe. Source: Enrichment cultures contain- ing acetate or butyrate as the only organic compound. Four strains were isolated from acetate enrichment cul- tures. The cultures were highly purified but not strictly pure. Habitat: Canal mud, sewage. Prob- ably occurs widely in fresh water sedi- ments where anaerobic conditions pre- vail. 2. Methanobacteritun omelianskii Bar- ker. (Bacille de la decomposition m6- thanique de I'alcohol ethylique, Omeliansky, Ann. Inst. Past., 30, 1916, 80; Barker, Arch f. Mikrobiol., 7, 1936, 436; also see Barker, Antonie van Leeu- wenhoek, 6, 1940, 201 and Jour. Biol. Chem., 137, 1941, 153.) Named for Prof. W. Omeliansky who first observed the organism. Rods: 0.6 to 0.7 by 1.5 to 10 microns, usual length 3 to 6 microns, unbranched, straight or slightly bent. Usually non- motile, occasionally feeble motility is observed. Spores of low heat resistance formed, spherical, terminal, swelling the rods. Primary alcohols, including ethyl, propyl, n-butyl and n-amyl alcohols, are oxidized to the corresponding fatty acids. Secondary alcohols, including isopropyl and sec-butyl, are oxidized to the corresponding ketones. Hydrogen is oxidized. Fatty and hydro.xy acids, glucose, polyalcohols and amino acids are not attacked. Carbon dioxide is used and converted to methane. Growth and alcohol oxida- tion are directly proportional to the carbon dioxide supply at low concen- trations. Nitrate, sulfate and oxygen cannot be used as oxidizing agents. Utilizes ethyl alcohol best of all or- ganic compounds. Utilizes ammonia as a nitrogen source. Growing range: pH 6.5 to 8.1. Optimum temperature 37° to 40°C. Maximum 46° to 48°C. Obligate anaerobe. Source: Soil, fresh water and marine muds, rabbit feces, sewage. Pure cul- tures were isolated from fresh water and marine muds (Barker, loc. cit., 1940). Habitat: Wherever organic matter is decomposing in an anaerobic, approxi- mately neutral environment. FAMILY BACTERIACEAE 647 Appendix III : Miscellaneous species of non-motile, or motile, non-spore-forming rod-shaped bacteria not previously listed or described. Ascobacterium luieurn Babes. (Babes, in Cornil and Babes, Les Bacteries, 3rd ed., 1, 1890, 155; also see Petri, Cent. f. Bakt., II Abt., 26, 1910, 359.) From water in Budapest (Babes) and the olive fly (Petri). Bacillus a, b, c, d, e,f, h and i, Vignal. (Arch. d. phys. norm, et path., Ser. 3, 8, 1886, 350-373; also see Flavobaderium biiccah's Bergey et al. and Bacillus bucca- lis fortuitus Sternberg.) From saliva and the teeth. Bacillus acido-aromaticus Van der Leek. (Cent. f. Bakt.. II Abt., 17, 1907, 652.) From milk. Bacillus acutangulus ^ligula. (Xo. 13, Lembke, Arch. f. Hyg., 29, 1897, 319; Migula, Syst. d. Bakt., 2, 1900, 680.) From feces. Bacillus acuttis Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 433.) From the stomach of a bird. Bacillus adavictzii jNIigula. (Bacillus Xo. XIII, Adametz, Landwirtsch. Jahrb., 18, 1889, 246; Migula, Syst. d. Bakt., 2, 1900, 686; not Bacillus adametzi Trevisan, I generi e Ic specie dello Batteriacee, 1889, 19.) From cheese. Bacillus aeris Chester. (Bacillus vio- laceus sacchari Ager, X. V. Med. Jour., 1894, 265; see Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 369; Bacterium violaceous sacchari Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 116; Chester, Man. Determ. Bact., 1901, 260.) From air. Produces a violaceus black pigment in old cultures in milk. Bacillus aerobius Doj'en. (Bacillus urinae aerobius Doyen, Jour. d. connaiss. medic, 1889, 107; Doyen, ibid.. 108.) From urine. Bacillus aerogencs Miller. (Miller, Deutsche med. Wchnschr., 12, 1886, 119; see Miller, Die ]\Iikroorganismen der jMundhohle, Leipzig, 1889, 262; not Ba- cillus aerogenes Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 340.) From the stomach. Bacillus aerogenes sputigenus capsula- tus Herla. (Arch, de Biol., 14, 1895, 403; abst. in Cent. f. Bakt., 25, 1899, 359.) From the blood of a mouse which had been inoculated with the sputum of a pneumonia patient. Bacillus aeschynomenus Trevisan. (Bacille de Pair j, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 150; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 20.) From air. Bacillus aethebius Trevisan. (Bacille de Pair c. Babes, in Cornil and Babes, Les Bacteries, 2nd cd.,1886, 149; Trevi- san, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus agilis Trevisan. (Bacillus der Vagus-Pneumonie, Schou, Fortschr. d. Medicin, 3, 1885, Xo. 15; Bacillus pneumonicus agilis Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 262; Trevisan, I generi e le specie delle Batteriacee, 1889, 14; not Bacillus agilis Tschisto- witsch, Berl. klin. Wochnschr., 1892, 512; not Bacillus agilis Chester, Man. De- term. Bact., 1901, 226; not Bacillus agilis ?klattes, Sitzungsber. d. Gesells. z. Beforderung d. gesam. Xaturw. z. Mar- burg, 62, 1927, 406; not Bacillus agilis Hauduroy et al., Diet. d. Bact. Path., Paris, 1937, 33; Bacterium pneumonicus agilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 140; Bacterixim vagus pneumonic Ch.Qster, ibid., 144; Ba- cillus pneumonicus Migula, Syst. ^\. Bakt., 2, 1900, 752.) From the lungs of rabbits having vagus pneumonia. Bacillus agtlis Mattes. (Sitzungsber. d. Gesells. z. Beforderung d. gesam. Xaturw.- z. :\Iarburg, 62, 1927, 406.) From the Meditenanean flour moth (Ephesiia kuehnielta). Bacillus agillimus DeToni and Trevi- san. (Bacillus lutcus pulidus Maggiora, Giorn. d. Soc. ital. d'Igiene, 11, 1889, 344; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, S, 1889, 969.) From the skin. 648 MAXUAL OF DETERMINATIVE BACTERIOLOGY Bacillus agnorum Trevisan. (Bacte- rium subtile agnorum Rivolta, Giorn. di Anat. fisiol. degli animali, 1881, 31 and 1883, 78; Trevisan, I generi e le specie delle Batteriacee, 1889, 13.) From dis- eased lambs. Bacillus alacer Eckstein. (Ztsclir. f. Forst- u. Jagdwesen, 26, 1894, 13.) Found associated with the eggs of the nun moth {Lymantria moyiacha). Bacillus alatus Grieg Smith. (Proc. Linn. Soc. New So. Wales, 30, 1905, 570.) Bacillus albatus Kern. . (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 408.) From the stomach and' intestines of a bird. Bacillus albiis Pagliani, Maggiora and Fratini. (Weisser Bacillus, Eisenberg, Bakt. Diag., 1 Aufi., 1886, Table 7; Pagliani et al., Giorn. d. Soc. ital. d'Igiene, 9, 1887, 587; not Bacillus albus Trevisan, 1 generi c le specie delle Batteriacee, 1889, 14; not Bacillus albus Bergey et al., Manual, 3rd ed., 1930, 398; Bacterium albus Chester, Ann. Kept. Del. Col. Agr. E.\p. Sta., 9, 1897, 76.) From water. Bacillus albus anaerobiescens Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 191.) From water. Bacillus albus putidus DeBar}^ (Quoted from Sternberg, Man. of Bact., 1893, 675.) From water. Bacillus albus- putidus Chester. (Ma- schek, see Adametz, Bakt. Nutz. u. Trinkwasser, 1888; Chester, Man. De- term. Bact., 1901, 237.) From water. Bacillus albus putridus Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 186.) From water. Bacillus alpha Dyar. (Ann. X. Y. Acad. Sci., 8, 1895, 366.) From air. Bacillus amabilis Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 358; Bacterium amabilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 110.) From air. Bacillus amarillae Trevisan. (Bacille de la fi^vre jaune. Babes, in Cornil and Babes, Les Bact^ries, 2nd ed , 1886, 529; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 13.) From a case of 3'^enow fever. Bacillus amarus Migula. (Bacillus liquefaciens lactis amari v. Freudenreich, Landwirlsch. Jahrb. d. Schweiz, 8, 1894; Migula, Syst. d. Bakt.,^, 1900, 694; not Bacillus amarus Hammer, Iowa Agr. Exp. Sta. Res. Bull. 52, 1919, 198.) From bitter milk. Bacillus amerimnus Trevisan. (Ba- cille de Fair b. Babes, in Cornil and Babes, Les Bact^ries, 2nd ed., 1886, 149; Trevisan, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus' amygdaloides Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 246.) From brine on salted pickles. Bacillus anceps Trevisan. (Bacille du mucus intestinal normal a. Babes, in Cornil and Babes, Les Bact^ries, 2nd ed., 1886, 153; Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) Fiom nor- mal intestinal mucous. Bacillus anthraciformis Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 28.) From meat extract. Bacillus anthracoides Trevisan. (Ba- cille de Pair /:, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 151; Trevisan, I generi e le specie delle Batteriacee, 1889, 20; not Bacillus an- thracoides Kruse, in Fliigge, Die jNIikro- organismen, 3 AuH., 2, 1896, 232.) From air. Bacillus annulatus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, 2, 1894, 30.) From water. Bacillus anularius Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 32.) From Emmenthal cheese. Bacillus apicum Kruse. (Canestrini, Atti Soc. Yen. Trent. Sci. Nat., 12, 1892, 134; Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufi., 2, 1896, 233.) From in- fected bees and their larvae. Bacillus apisepticus Burnside. (Jour. Econ. Ent., 21, 1928, 379.) Pathogenic for the honey bee (Apis mellifera). Bacillus aquatilis Migula. (Bacillus FAMILY BACTERIACEAE 649 nquatilis sulcatus IV, Weichselbaum, Das osterreifhischo Sanitiitswesen, 1889, No. 14-23; Bacterium uquatilis sulcatus quartus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 72; Migiila, Syst. d. Bakt., 2, 1900, 733; not Bacillus aqua- tilts Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 381; Bacillus aquatilis- sidcatus-quartus Chester, Man. Determ. Bact., 1901, 216.) From water. Bacillus aquatilis Trevisan. (Bacille de I'eau a. Babes, in Cornil and Babes, Les Bac t eries , 2nd ed . , 1886 , 167 ; Trevisan , I generi c le specie delle Batteriacee, 1889, 19; not Bacillus aquatilis Frank- land and Frankland, Ztschr. f. Hyg., 6', 1889,381.) From water. Bacillus aquatilis conununis Kruse. (Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., £, 1896. 315; Bacterium aquatilis communis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 91.) Found commonly in water. Listed bj^ Kruse as a non-chromogenic strain of Bacillus fluorescens liquefaciens {Pseudomonas jluorescens Migula). Bacillus arborescens Jamieson and Edington. (Brit. Med. Jour., 1, 1887, 1265.) From the desquamation of scar- let fever patients. Bacillus arboreus Migula. (Biium- chenbacillus, Maschek, Bakt. Unter- such. d. Leitmeritzer Trinkwasser, Leit- meritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 710.) From water. Bacillus aromaticus Beijerinck. (Quoted from Van der Leek, Cent. f. Bakt., II Abt., 17, 1907, 490; not Bacillus aromaticus Pammel, Bull. 20, Iowa Agr. Exp. Sta., 1893, 792; not Bacillus aro- maticus Grimm, Cent. f. Bakt., II Abt., 8, 1902, 589; not Bacillus aromaticus Van der Leek, loc. cit., 659.) From milk. Bacillus aromaticus Van der Leek. (Van der Leek, Cent. f. Bakt., II Abt., 17, 1907, 659.) From soft cheeses. Bacillus assimilis Trevisan. (Bacille de Fair i. Babes, in Cornil and Babes, Les Bact^ries, 2nd ed., 1886, 150; Trevisan, I generi e le specie delle Batteriacee, 1899, 20.) From air. Bacillus aurantius Trevisan. (Or- angerother Wasserbacillus, Adametz and Wichmann, Mitth. Oest. \'ers. Stat. f. Brauerei u. Malz. in Wien, 1, 1888, 50; Trevisan, I generi e le specie delle Batteriacee, 1889, 19; not Bacillus aurayi- tius Bergey et al.. Manual, 3rd ed., 1930, 421.) From water. Bacillus aureus Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 9; prob- ablj- not Bacillus aureus Frankland and Frankland, Philos. Trans. Roy. Soc. London, 178, B, 1887, 272 and probably not Bacillus aureus Pansini, Arch. f. path. Anat. u. Physiol., 122, 1890, 436; not Bacillus aureus Adametz, quoted from Sternberg, Man. of Bact., 1893, 621.) Capable of infecting the larvae of various insects. Bacillus azureus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, 2, 1894, 24.) From water. Bacillus babesi Trevisan. (Bacille du mucus intestinal normal b. Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 153; Trevisan. I generi e le specie delle Batteriacee, 1889, 15.) From nor- mal intestinal mucus. Bacillus belfa7itii Migula. (Eine neue pathogene Bakteriumart im Tetanus- material, Belfanti and Pescarolo, Cent. f. Bakt., 4, 1888, 51Z; Bacillus accidentalis tetoni Kruse, in Fliigge, Die ]\Iikro- organismen, 3 Aufl., 2, 1896, 433; Bacte- rium accidentalis tetani Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 88; Migula, Syst. d. Bakt., 2, 1900, 767; not Bacillus belfanti Carbone and Ven- turelli. Boll. 1st. Sieroter., Milan, 4, 1925, 59; Bacillus accidentalis Chester, Man. Determ. Bact., 1901, 229.) From pus in a case of tetanus. Bacillus benzoli Tausson. (Planta, 7, 1929, 735.) From soil. Oxidizes ben- zene. Bacillus beribericus Trevisan. (Ca- ratt. di ale. nuov. gen. di Batt., 1885, 12.) From cases of beri-beri in Japan, G50 MAiNTUAL OF DETERMINATIVE BACTERIOLOGY Also see Ogata, abst . in Cent. f. Bakt., 3, 1888, 75. Bacillus berolinensis Migula. (Uutci- Bacillus aus Wasser, P'raenkel, Grundriss der Bakterienkunde, 3 Aufl., 1890, 252; Bacillus ruber berolinerxsis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 303; Bacterium ruber berolinensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 113; Migula, Syst. d. Bakt., 2, 1900, 856; not Bacillus berolin- ensis Chester, Man. Determ. Bact., 1901, 305.) From water. Rust-red to orange-yellow pigment on potato. Bacillus beta Dyar. (Ann. X. Y. Acad. Sci., 8, 1895, 366.) From air. Bacillus beyerinckii DeToni and Trevi- san. (Bacillus radiciculu var. lique- facicns Beijerinck, Bot. Zeitung, 1888, 750; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 972; not Ba- cillus beijerinckii Henneberg, Ztschr. f. Spiritusindustrie, 26, 1903, 22; see Cent, f. Bakt., II Abt., 11, 1903, 159.) From soil and the roots of legumes. Bacillus billim/si Chester. (Bacillus of corn-stalk disease of cattle, Billings, in Baumgarten, Jahresbericht, 1889, 184; Chester, .Man. Determ. Bact., 1901, 214.) Isolated bj^ liillings from corn-stalk disease of cattle, and by Xocard from bronchopneumonia in oxen. Bacillus bombycis Chatton. (Chatton, Compt. rend. Acad. Sci., Paris, 156, 1913, 1708; not Bacillus bombycis Macchiati, Stazioni sperimentali Agrarie Italiane, 20, 1891, V2l; Bacterium bombycis Paillot, L'infection chez les insectes, 1933, 131.) From diseased silkworms (Bombyx mori). Bacillus bookeri Dj'ar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 378; not Bacillus bookeri Ford, Studies from the Royal Victoria Hospital, Montreal, /, 1903, 31.) Found by Dr. I'ruddeii in a case of cj's- litis. Bacillus bracliythrix DeToni and Trevi- san. (Bacillus G, Maggiora, Giorn. Soc. ital. d'Igiene, 11, 1889, 348; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 967.) From the skin. Bacillus brunneus (Schroeter) Schroc- ter. (Bacteridium brunneum Schroetei', in Cohn, Beitr. z. Biol. d. Pflanz., /, Heft 2, 1872, 126; Schroeter, in Cohn, Krj'ptog. Flora v. Schlesien, 3 (1), 1886, 158; not Bacillus brunneus Adametz and Wichmann, Die Bakt. der Nutz- und Trinkwasser, Wien, 1888; Bacillus fuscus Fliigge, Die Mikroorganismen, 2 Aufi., 1886, 290; not Bacillus fuscus Zimmer- mann, Bakt. unserer Trink- u. Nutz- wjisser, Chemnitz, 1, 1890, 70.) Bac- terium brunneum Schroeter or Cohn is given as a synonym by Fliigge (1886) and by Trevisan (1889) but this appears to be an incorrect spelling of Bacteridium brunneum Schroeter. Neither Scliroeter nor Cohn used Bacterium brunneum in 1872 or later so far as can be determined by a careful study of their papers. From corn, wheat and potato infusions. Bacillus huccalis fortuitus Sternberg. (Bacillus j, Vignal, Arch. Phys. norm, et path., S^r. 3, 8, 1886, 337; Sternberg, Man. of Bact., 1893, 685; Bacterium bu- calis (sic) fortuitus Chester, Ann. Rept. Del. Col. Agr. E.\p. Sta., 9, 1897, 91 and 130; Bacillus ttucalis (sic) Chester, Man. Determ. Bact., 1901, 234; not Bacillus buccalis Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) From the mouth. Bacillus buccalis mucifercns Miller. (Miller, Dental Cosmos, 33, 1891, 792 and 800.) From the blood. A slimy capsulated bacillus. Bacillus buccalis septicus Miller. (Miller, Dental Cosmos, 33, 1891, 792 and 802.) From the mouth and in pus of an abscess caused bj^ a dental instrument. Bacillus butyri Migula. (Bacillus bu- tyri I, V. Klecki, Cent. f. Bakt., 15, 1894, 357; Migula, Syst. d. Bakt., 2, 1900, v and 811.) From rancid butter. Bacillus caeci Ford. (Studies from the Royal Victoria Hosp., Montreal, 1, (5), 1903, 45; also see Jour. Med. Res., FAMILY BACTERIACEAE 651 1, 1901, 217.) From the stomach and rectum. Bacillus canalensis Castcllani. (Proc. Soc. Exp. Biol, and Med., 25, 1928, 540.) From human feces. Bacillus canvs Migula. (Grauer Ba- cillus, Masehek, Untersuch. d. Leitmerit- zer Trinkwasser, Leitmeritz, 1887; Mi- gula, Syst. d. Bakl., .?, 1900,711.) From water. Bacillus canus Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 24, 1894, 15.) From larvae of the nun moth {Lymantria monacha.) Bacillus carabijormis Raczyscki. (Diss, milit. medic. Acad. Petropolitanae Ruteniae, 1888; abst. in Cent. f. Bakt., 6, 1889, 113.) From the stomach of a dog. Bacillus carnis Wilhelmj'. (Wilhelmj-, Arb. bakt. Inst. Karlsruhe, 3, 1903, 21; not Bacillus camis Klein, Cent. f. Bakt., II Abt., 35, 1903, 459.) From meat extract. Bacillus caseolyticus Lochmann. (Cent. f. Bakt., I Abt., Orig., 31, 1902, 385.) From the organs of guinea pigs which had been inoculated with tubercle bacilli. Bacillus cathetus Trevisan. (Bacille de Fair g. Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 150; Trevi- san, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus caviae Trevisan. (Bacille du mucus intestinal normal du cobaye e, Babes, in Cornil and Babes, Les Bac- teries, 2nd ed., 1886, 154; Trevisan, I generi e le specie delle Batteriacee, 1889, 15; Pasteurella caviae DeToni and Trevi- san, in Saccardo, Sylloge Fungorum, 8, 1889, 996; not Pasteurella caviae Ilaudu- roy et al.. Diet. d. Bact. Path., 1937, 313.) From the intestinal mucus of guinea pigs. Bacillus centralis Zimmermann. (Bakt. unserer Trink. u. Nutzwasser, Chemnitz, 2, 1894, 10.) From water. Bacillus charrini Trevisan. (Bacille de la pseudo-tuberculosc bacillaire du cobaye, Charrin and Roger, Compt. rend. Acad. Sci., Paris, 106, 1888. 868; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 13.) From pulmonary tuberculosis of guinea pigs. Bacillus chlorinus Migula. (Griingcl- ber Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 50; Migula, Syst. d. Bakt., 2, 1900, 820; not Bacillus chlorinus Frank- land and Frankland, Philos. Trans. Roy. Soc. London, 178, B, 1887, 274.) From water. Bacillus chyluriae Trevisan. (Bacil- lus of chyluria, Wilson, Brit. Med. Jour., No. 1249, 1884, 1128; Trevisan, Atti Acad. Med. -Fis. -Stat. Milan., Ser. V, 3, 1885, 99.) From chyluria. Bacillus citreus (Unna and Tommasoli) Kruse. {AscohacJllus cilreus Unna and Tommasoli, Monats. f. prakt. Dermatol., 9, 1890, 60; Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 309; not Bacillus citreus Frankland and Frank- land, Philos. Trans. Roy. Soc, London, 178, 1887, B, 272; Bacterium citreus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 104.) From the human skin in cases of eczema. Bacillus citricus Kern. (Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 426; not Bacillus citricus Weiss, ibid., 2, Heft 2, 1902, 234.) From the intes- tines of birds. Bacillus citrnius Migula. (Citrongel- ber Bacillus, Masehek, Bakt. I'ntersuch. d. Leitmeritzer Trinkwasser, 1887; Mi- gula, Sj-st. d. Bakt., 2, 1900, 832.) From water. Bacillus cladogenes Trevisan. (Bac- terie de Fair No. 3, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 140; Trevisan, I generi e le specie delle Batteri-acee, 1889, 20.) From air. Bacillus claviformis Doyen. {Bacillus urinae claviformis Doyen, Jour. d. con- naiss. medic, 1889, 106; Doyen, ibid., 108; Bacillus doyeni DeToni and Trevi- 652 MANUAL OF DETERMINATIVE BACTERIOLOGY sail, in Saccardo, Sylloge Fungorum, 8, 1889, 949.) From urine. Bacillus cleoni Picard. (Bull. Soc. d'Etude et de Vulgarisation Zool. Agric, 13, 1913, 134.) A fluorescent cocco- bacillus. From diseased larvae of weevils {Temnorrhinus (Cleonus) mendi- cus). Bacillus coccineus Catiano. (Catiano, in Cohn, Beitr. z. Biol. d. Pflanz., 7, 1896, 339; not Bacillus coccineus Pansini, Arch. f. path. Anat., 122, 1890, 437; Bacillus subcoccineus Migula, Syst. d. Bakt., 2, 1900, 857.) From the vagina. Reddish pigment. Bacillus coeruleo-viridis Trevisan. (Blaugriin fluorescirende Bacterium, Adametz, IMitth. Oest. Vers. Stat. f. Brauerei u. Malz. in Wien, 1, 1888, 46; Trevisan, I generi e le specie dellc Bat- teriacee, 1889, 20; Bacterium coeruleo- viride DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1087.) From water. Bacilhis coeruleus Eckstein. (Eck- stein, Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 14; not Bacillus coeruleus Smith, Med. News, 1887, 758; probably not Bacillus coeruleus Voges, Cent. f. Bakt., 14, 1893, 301.) From larvae of the nun moth (Lymantria monacha). Bacillus columbarum Chester. (Ba- cillus of pigeon cholera, Moore, U.S. D.A., Bur. Anim. Ind., Bull. 8, 1895; Chester, Man. Determ. Bact., 1901, 209.) From a disease of pigeons. Had- ley et al. (Rhode Island Agr. Exp. Sta., Bull. 174, 1918, 178) regard this as prob- ably a paracolon. Bacillus constricius Zimmermann. (Zimmermann, Bakt. unserer Trink- u. Nutzwasser, Chemnitz, 1, 1890, 42; Bacterium constricius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 112.) From water. Bacillus convolutus Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 461; Bacterium convolutus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 101.) From river water. Bacillus coprogenes foetidus Sternberg. (Darmbacillus, Schottelius, 1885; Stern- berg, Man. of Bact., 1893, 468.) From the intestinal contents of pigs which had died of swine erysipelas. Bacillus coroncitus Keck. (Inaug. Diss., Dorpat, 1890, 43.) From water. Bncillus corvi Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 394.) From the stomach and intestines of birds. Bacillus courmontii Migula. (Cour- mont, Compt. rend. Soc. Biol., Paris, 1889; Bacillus pseudotuberculosis si7nilis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 454; Migula, Syst. d. Bakt., 2, 1900, 770.) From tubercles of cattle. Bacillus crassus Lucet. {Bacillus crassus pyocjenes bovis Lucet, Ann. Inst. Past., 7, 1897, 327; Bacillus crassus pyo- genes Lucet, ibid., 327; Lucet, ibid., 328; Bacillus pyogenes crassus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 343; Bacterium pyogenes crassus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 141; Bacillus bovis Migula, Syst. d. Bakt., 2, 1900, 765.) From bo- vine abscesses. Regarded by Kruse as a synonym of Bacillus pneumoniae. Bacillus crinitus Migula. (No. 15, Lembke, Arch. f. Hyg., 29, 1897, 321; Migula, Syst. d. Bakt., 2, 1900, 678.) From feces. Bacillus cubonianus Cuboni and Gar- bini. (Atti. dei Lincei, Ser. 4, 6, 1890, 26-27, ciuoted from Steinhaus, Bact. Assoc. Extracell. with Insects and Ticks, Minneapolis, 1942, 53; not Bacillus cu- bonianus Macchiati, Staz. Sperim. Agr. Ital., 23, 1892, 228.) From silkworms {Bombyx mori). Bacillus cuenoti Mercier. (Bakteri- enjihnlichen Gebilden, Blochmann, Ztschr. f. Biol., 24, 1887, 1; Compt. rend. Soc. Biol., Paris, 61, 1906, 682; also in Arch. f. Protistenkunde, 9, 1907, 346.) From the fat body of the cockroach (Periplaneta orientalis) . Bacillus cuniculi Migula. (Bacillus septicus cuniculi Lucet, Ann. Inst. Past., 6", 1892, 564; Bacillus cuniculi septicus Kruse, in Fliigge, Die Mikroorganismen, FAMILY HACTERIACEAE (i53 3 Aufl., 2, 1896, 406; Bacterium cunicuH septicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 76; Migula, Syst. d. Bakt., 2, 1900, 758.) Associated with a spontaneous epizootic of rabbits. Bacillus cunicnlicida immohilis Kruse. (Kruse, in P^liigge, Die Mikroorganis- men, 3 Aufl.. ^, 1896, 417; Bacterium cuniculicida immobilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, SA; Bacterium cuniculicida var. immobile Chester, Man. Determ. Bact., 1901, 140.) Cause of a disease of rabbits. Bacillus cystiformis Clado. (Quoted from Sternberg, Man. of Bact., 1893, 649). From urine in a case of cystitis. Bacillus cystitidis Migula. (Cocco- bacillus aerogenes vesicae Schow. Cent. f. Bakt., 1^, 1892, 749; Bacillus aerogenes ■vesicae Lehmann and Neumann, Bakt. Diag., 1 Aufl., i", 1896, 237; Bacterium aerogenes vesicae Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 128; Migula, Syst. d. Bakt., 2, 1900, 771; Bacillus aerogenes Chester, Man. De- term. Bact., 1901, 227; not Bacillus aero- genes Miller, Deutsche med. Wochnschr., 12, 1886, 119; not Bacillus aerogenes Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufl., £, 1896, 340.) From urine in a case of cystitis. Bacillus dacryoideus Migula. {Bacil- lus oogenes hydrosulfureus rj, Zorken- dorfer. Arch. f. Hyg.. 16. 1893, 389; Migula, Syst. d. Bakt.. 2. 1900, 791.) From hens' eggs. Bacillus decolor Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26. 1894, 15.) From the larvae of a butterfly {Vanessa utricae). Bacillus decolorans major Dyai'. (Ann. X. Y. Acad. Sci., 8. 1895, 362.) From air. Bacillus decolorans minor Dyar. (Ann. N. Y. Acad. Sci., 8. 1895, 359.) From air. Bacillus defessus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 397.) From the stomach and intestines of birds. Bacillus delta Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 368; Bacterium delta Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., .9, 1897, 114.) From water. Bacillus denitrificans Giltay and Aber- son. (Gilta,y and Aberson, Arch. Neerl. Sci. exact, et nat., 2S, 1891, 341; quoted from Sternberg, Man. of Bact., 1893, 727; not Bacillus denitrificans Migula, Syst. d. Bakt., 2, 1900, 796; not Bacillus denitrificans Chester, Man. Determ. Bact., 1901, 274.) From soil and air. Bacillus deintrofluorescens van Iterson. (Cent. f. Bakt., II Abt., .9, 1902, 772; 12, 1904, 111.) Fluorescent. From soil. Bacillus dentalis viridans Miller. (Miller, Die Mikroorganismen der Mund- hohle, Leipzig, 1889, 218.) From carious teeth. Bacillus dermoides Tataroff. (Inaug. Diss., Dorpat, 1891, 19.) From water. Bacillus diaphanus Migula. (Hali- bacterium pellucidum Fischer, Die Bak- terien des Meeres, 1894, 22; Migula, Syst. (1. Bakt., 2, 1900, 712.) From sea water. Bacillus diffluens Doyen. {Bacillus urinae difiluens Doyen, Jour. d. connaiss. medic, 1889, 107; Doyen, ibid., 108; not Bacillus difiluens Castellani,. 1915, see Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 943.) From urine. Bacillus digitatus Migula. (Bacillus No. 7, Pansini, Arch. f. i)ath. Anat., 122, 1890, 443; Migula, Syst. d. Bakt., 2, 1900, 659.) From sputum. Bacillus dissimilis Trevisan. (Bacil- lus I, Leube, Arch. f. path. Anat., 100, 1SS5, 556; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From urine. Bacillus domesticus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 358; Bacterium domesticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, no.) From air. Bacillus droserae (Troili-Petersson) Buchanan and Hammer. {Bacterium droserae Troili-Petersson, Cent. f. Bakt., II Abt., 38, 1913, 1; Buchanan and Ham- mer, Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 256.) Isolated by placing leaves of a sundew {Drosera. intermedia) in milk and isolating the 654 MANUAL OF DETERMINATIVE BACTERIOLOGY slimy milk organisms developing. Closely related to Bacterium lacto- rubefaciens Gruber, according to Bu- chanan and Hammer. Bacillus duclauxii (Miquel) DeToni and Trevisan. (Urobacillus duclauxii Miquel, Ann. d. Microgr., 2, 1889, 58; DeToni and Trevisan, in Saccardo, Syl- loge Fungorum, 8, 1889, 963.) From sewage . Bacillus eczemicus Trevisan. (I generi e le specie delle Batteriacee, 1889, 14.) From exudate in cases of eczema. Bacillus egregius Zopf. (Quoted from Papenhausen, Arb. bakt. Inst. Karls- ruhe, 3, 1903, 59.) A reddish-yellow non-spore-forming rod. Bacillus elipsoideus Migula. {Bacillus saprogenes vini v, Kramer, Die Bak- terien in ihren Beziehungen zur Land- wirtschaft, 2, 1892, 138; Migula, Syst. d. Bakt., 2, 1900, 684.) From wine. Bacillus ellingtonii Chester. (Bacil- lus No. 21, Conn, Rept. Conn. (Storrs) Agr. Exp. Sta., 1893, 52; Chester, Man. Determ. Bact., 1901, 264.) From milk. Bacillus emmans Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 232.) From vegetable infusions. Bacillus emulsinus Fermi and Monte- sano. (Cent. f. Bakt., 15, 1894, 722.) From air. Decomposes amygdalin. Bacillus endocarditidis Migula. (Ba- cillus endocarditidis griseus Weichsel- baum, Beitr. z. path. Anat., 4, 1889, 119; Bacterium endocarditidis griseus Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 88; Migula, Syst. d. Bakt., 2, 1900, 750.) From a case of endocarditis. Bacillus engelmanni Trevisan. (Bac- terium chlorinum Engelmann, see Flugge, Die Mikroorganismen, 2 Aufl., 1886, 289; not Bacterium chlorinum Migula, Syst. d. Bakt., 2, 1900, 471; Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) Source not given. Bacillus enleromijces Trevisan. (Ba- cille des selles /, Babes, in Cornil and Babes, Les Bact^ries, 2nd ed., 1886, 154; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 15.) From feces. Bacillus entomotoxicon Duggar. (Bull. Illinois State Lab. Nat. Hist., 4, 1896, 340-379.) From the squash bug (Anasa tristis) . Bacillus epsilon Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 369; Bacterium epsilon Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 114.) From air. Bacillus equi Migula. (Bacillus equi intestinalis Dyar and Keith, Technol. Quarterly, 6, 1893, No. 3; abst. in Cent. f. Bakt., 16, 1894, 838; Bacterium equi in- testinalis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 70; Migula, Syst. d. Bakt., 2, 1900, 87 i; Bacillus intestinalis Chester, Man. Determ. Bact., 1901, 213.) From the intestines of a horse. Bacillus erubescens Migula. (Bacillus oogenes hydros ulfureus k, Zorkendorfer, Arch. f. Hyg., 16, 1893, 391; Migula, Syst. d. Bakt., 2, 1900, 792; Bacillus rubescens Nepveux, These, Fac. Pharm. Paris, 1920, 113.) From hens' eggs. Bacillus erijthrogenes rugatus Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 374.) A wrinkled variety of Bacillus lactis erythrogenes Hueppe. Bacillus erythrosporus Miflet. (Mifiet, in Cohn, Beitr. z. Biol. d. Pflanz., S, Heft 1, 1879, 135; Bacillus (Streptobacter) erylhrospores (sic) Schroeter, in Cohn, Krypt. Flora v. Schles., 3, 1, 1886, 158; Bacterium erythrosporus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 123.) From putrefying egg-white and meat infusion. According to Chester, the author mistook reddish granules for spores. Fluorescent. Bacillus esterificans fluorescens Maa- sen. (Arb. a. d. k. Gesundsheitsamte, 15, 1899, 504-507.) From grains and from rotting vegetation in river water. Bacterium esterificans stralauense Maa- sen. (Arb. a. d. k. Gesundheitsamte, 15, 1899, 504-507.) From Spree River water. Bacillus eta Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 374; Bacterium eta Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 107.) From air. Bacillus ethareticus Frankland and FARHLY BACTERIACEAE 655 Frankland. (Proc. Roy. Soc. London, 46, 1889.) Ferments mannitol, glycerol and glucose to ethyl alcohol and acetic acid with a trace of formic and succinic acids. Bacillus ethacelosuccinicus Frankland and Frew. (Transactions of the Chemi- cal Society, 1892, 275.) Ferments man- nitol and dulcitol to ethyl alcohol, acetic acid, succinic acid, hydrogen and car- bonic acid. Bacillus exapatus Trevisan. (Bacillus der conjunctivalsack /, Fick, Ueber Mikroorg. in Conjunctivalsack, Wies- baden, 1887; Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) Found frequently in the human eye. Bacillus exiguus Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 447; Bacterium exiguum Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 114; not Bacterium exiguum Staubli, Miinch- ner med. Wochnschr., Xo. 45, 1905.) From water. Bacillus faniigcr Trevisan. (Bacillus bei Erysipel am Kaninchenohr, Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 283; Trevisan, I generi e le specie delle Batteriacee, 1889, 14.) From a case of erysipelas of the ear of a rabbit. Bacillus fclis (Rivolta) Trevisan. {Cocco-tacterium felis Rivolta, Giorn. di Anatomia, No. 1, 1888; Trevisan, 1 generi e le specie delle Batteriacee, 1889, 14.) From an infection in a cat. Bacillus fermentationis Chester. (Ba- cillus foetidus liquefaciens Tavel, Ueber d. Aetiol. d. Strumitis, Basel, 1892; Chester, Man. Determ. Bact., 1901, 233.) From strumitis. Bacillus ferrugineus Rullmann. (Rullmann, Cent. f. Bakt., I Abt., 24, 1898, 467; not Bacillus ferrugineus Van Iterson, Cent. f. Bakt ., II Abt., 11, 1903, 694.) From canal water. Bacillus ferrugineus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 361; not Bacillus ferrugineus Rullmann, Cent. f. Bakt., I Abt., Orig., U, 1898, 465; Bac- terium furrugineus (sic) Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 115; Bacterium ferrugineum Chester, Man. Determ. Bact., 1901, 177.) From air and from a fresh leaf of the pitcher plant (Sarracenia purpurea) . Bacillus feriilis DeToni and Trevisan. (Bacillus urinae feriilis Doyen, Jour. d. connaiss. medic, 1889, 107; DeToni and Trevisan, in Saccardo, Sylloge Fungo- rum, 8, 1889, 949.) From urine. Bacillus figurans Vaughan. (Vaughan, Amer. Jour. Med. Sci., 104, 1892, 107; not Bacillus figurans Crookshank, Man. of Bact., 1st ed., 1886.) From water. Bacillus finitimus ruber Dyar. (Ann. N.Y. Acad. Sci., S, 1895,361.) From air. Bacillus flavidescens ^Sligula. (Ba- cillus aquatilis sulcatus v, Weichsel- baum, Das osterreichische Sanitatswe- sen, 1889, No. 14-23; Bacillus aquatilis sulcatus Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 382; Migula, Syst. d. Bakt., 2, 1900, 734; Bacillus weichselbaumii Chester, Man. Determ. Bact., 1901, 218.) From water. Bacillus flavoides Castellani. (Proc. Soc. Exp. Biol, and :\Ied., 2S, 1928, 539.) From the human skin. Bacillus flavus Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 12; not Bacillus flavus Fuhrmann, Cent. f. Bakt., II Abt., 19, 1907, 117; not Bacillus flavus Bergey et al.. Manual, 1st ed., 1923, 286.) From dead larvae of a butterflj^ (Vanessa polychlorus). Bacillus flexuosus Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 460; Bacterium flexuosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 100.) From river water. Bacillus floccosus Kern. (Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 424; not Bacillus floccosus Weinberg et al., Les Microbes Anaerobies, 1937, 698.) From the stomach and intestines of birds. Bacillus flu idificans DeToni and Trevi- san. (Bacillus fluidificans parvus iNIag- giora, Giorn. Soc. ital. d'Igiene, 11, 1889, 344; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 969.) From the skin. 656 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacillus fluorescens Chester. (Bacil- lus fluorescens aureus Zimmermann, Bakt. unserer Trink- u. Xutzwjisser, Chemnitz, 1, 1890, 14; Bacterium fluo- rescens aureus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 109; Chester, Man. Determ. Bact., 1901, 255; not Ba- cillus fluorescens Trevisan, I generi e le specie delle Batteriacee, 1889, 19; not Bacillus fluorescens Bergey et al., Man- ual, 1st ed., 1923, 287.) From water. Bacillus fluorescens Pagliani, Maggiora and Fratini. (Pagliani et al., Giorn. d. Soc. ital. d'Igiene, 9, 1887, 587; not Ba- cillus fluoresceiis Bergey et al., Manual. 1st ed., 1923, 287; Bacillus pacjlianii Trevisan, I generi e Ic specie delle Batteriacee, 1889, 19; not Bacillus pay- lianii Carbone and Venturelli, Boll. 1st. Sieroter., Milan, 4, 1925, 59.) From water and soil. Bacillus fluorescens alhus Zimmer- mann. (Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, /, 1890, LS.) From water. Bacillus fluorescens baregensis Robine and Hauduroy. (Compt. rend. Soc. Biol. Paris, 98, 1928, 25.) From the water of hot sulfur springs. Fourment (Compt. rend. Soc. Biol. Paris, 98, 1928, 588) states that this organism is a variety of Bacillus fluorescens liquefaciens Kruse, but Robine and Hauduroy (Compt. rend. Soc. Biol. Paris, 99, 1928, 317j deny this. Bacillus fluorescens liquefaciens minu- lissimus Unna and Tommasoli . (Quoted from Sternberg, Man. of Bact., 1893, 636.) From the surface of the body in cases of eczema seborrhoeicum. Bacillus fluorescens lonyus Zimmer- mann. (Zinmiermann, Bakt. unserer Trink- u. Nutzwiisser, Chemnitz, 1, 1890, 20; Bacterium fluorescens longus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124.) From water. Bacillus fluorescens tenuis Zimmer- mann. (Zimmermann, Bakt. unserer Trink- u. NutzwJisser, Chemnitz, 1, 1890, 14; Bacterium fluorescens tenuis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124.) From water. Bacillus foetidus Eckstein. (Ztschr. f. Forst- u.. Jagdwesen, 26, 1894, 12.) From dead larvae of a butterfly (Fanesso urlicae) . Bacillus foetidissimus Migula. (Ba- cillus pyogenes foetidus liquefaciens Lanz, Cent. f. Bakt., U, 1893, 269; Bacterium pyogenes foetidus liquefaciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 92; Migula, Syst. d. Bakt., 2, 1900, 643.) From a brain abscess. Bacillus fortissimus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 209.) From asparagus and brewer's grain infusions. Bacillus fuliginosus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 224.) From vegetable infusions. Bacillus fulvus Edington. (Edington, Brit. xMed. Jour., June 11, 1887, 1262; not Bacillus fulvus Zimmermann, Bakt. unserer Trink- u. Nutzwasser. Chem- nitz, /, 1890, 44; not Bacillus fulvus Migula, Syst. d. Bakt., 2, 1900, 844.) Associated with cases of scarlatina. Not pathogenic. Bacillus fulvus Migula. (Bacillus tu- berigenus 4, Gonnermann, Landwirtsch. Jahrb., 23, 1894, 656; Migula, Syst. d. Bakt., 2, 1900, 844; not Bacillus fulvus Zimmermann, Bakt. unserer Trink- u. Nutzwasser, Chemnitz, /, 1890, 44.) From root nodules on lupine. Bacillus fumeus Migula. (No. 5, Lembke, Arch. f. Hyg., 29, 1897, 313; Migula, Syst. d. Bakt., 3, 1900, 787.) From feces. Bacillus fumosus Migula. (No. 4, Lembke. Arch. f. Hyg., 29, 1897, 312; Migula, Syst. d. Bakt., 2, 1900, 788.) From feces. Bacillus fungosus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3. 1902, 255.) From fermenting beets. Bacillus fuscans Miller. (Miller, Die Mikroorganismen der Mundhohle, Leip- zig, 1889, 70.) From the mouth. Bacillus fuscescens Migula. (Bacillus fuscus limhatus Scheibenzuber, Allgem. Wiener med. Zeitung, 3J,, 1889, 171; FAMILY BACTERIACEAE 657 Bnrterinni fiiscus linihatus Chester. Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 117; Migula, Syst. d. Bakt., 3, 1900, 836; Bacterium fuscescens Chester, Man. Determ. Bact., 1901, 179; Bacillus fuscus Chester, ibid., 261; not Bacillus fuscus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 290; not Bacillus fuscus Zimmer- mann, Bakt. unserer Trink- u. Xutz- wasser, Chemnitz, 1, 1890, 70.) From rotten eggs. Bacillus galtieri Trevisan. (Microbe pathogone chromo-aromatique du pore, Galtier, Compt. rend. Acad. Sci. Paris, 106, 1888, 1368; Trevisan, I generi e le specie delle Batteriaoee, 1889, 14; Ba- cillus chromnaromalicus Eisenberg, Bakt. Diag., 3 Aufl., 1891^ 360; Bacterium chro- mo-aromaticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 119; Bacillus helvolns Chester, Man. Determ. Bact., 1901, 264; not Bacillus helvolus Zimmer- mann, Bakt. unserer Trink- u. Xutz- wasser, Chemnitz, 1, 1890, 52.) From lesions in a ease of broncho-pneumonia in hogs. Bacillus (jasoformans Pribram. {Bac- terium aquatile gasofor7na7is non liquc- faciens von Rigler, Hyg. Rund., 12, 1902, 482; Bacterium (jasoformans non lique- faciens von Rigler, ibid., 485 and Cent, f. Bakt., I Abt., Ref., 31, 1902, 682; Bacillus aquatilis gasoformans non lique- faciens Pribram, Klassification der Schi- zomyceten, Leipzig und Wien, 1933, 83; ibid., 83.) From bottled mineral waters. Similar to coliform bacteria e.xcept that it is a yellow chromogen. Bacillus gaytoni Cheshire. (Bees and Bee Keeping, London, 2, Part 13, 1886, 543 and 569.) Found in black bees (Afis mellifera), i.e., black because of the loss of hairy covering. Bacillus gelatinosus Migula. (Bac- terium gelatinosum betac Glaser, Cent, f. Bakt., II Abt., /, 1895, 879; Migula, Syst. d. Bakt.. 2. 1900. 805.) From beet juice. Bacillus gclatogenes Black. (Trans. 111. State Dental Soc. 22, 1886, 187.) From the mouth. Bacillus geton Trevisan. (Bacille de I'eau b, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 168; Trevisan, I generi e le specie delle Batteriacee, 1889, 19.) From water. Bacillus gigas Goot. (Goot, Med. Proefstat. voor de Java Suikerindustrie, Pt. 5, Xo. 10, 60 pp., quoted from Stein- haus, Bact. Assoc. Extracell. with In- sects and Ticks, Minneapolis, 1942, 58; not Bacillus gigas Zeissler and Rasse- feld. Arch. f. wiss. u. prakt. Tierheilk., 59, 1929, 419.) From larval and adult stages of a beetle (Adoretus cotnpressus) . Bacillus (/) gingivae -Migula. (Bac- terium gingivae pyogenes IMiller, Die Mikroorganismen der iVIundhohle, Leip- zig, 1889, 217; Bacillus pyogenes gingivae Kruse, in Fliigge, Die ^likroorganismen, 3 Aufl., 2, 1896, 287; Migula, Syst. d. Bakt., 2, 1900, 642.) From the oral cavity. Bacillus glaucus Maschek. (Maschek, Bakt. I^ntersuch. d. Leitmeritz. Trink- wasser, Jahresber. d. Oberrealschule zu Leitmeritz, 1887; quoted from Sternberg, Man. of Bact., 1893, 637; Bacterium glau- cus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 116.) From water. Bacillus globosus Migula. (Bacterium .4, Peters, Botan. Zeit., 47, 1889; Migula, Syst. d. Bakt., 2, 1900, 798.) From fer- menting dough. Bacillus globulosus Weiss. (Arb. bakt . Inst. Karlsruhe, 2, Heft 3, 1902, 253.) From sauerkraut and bean infusions. Bacillus gonnermanni Migula. {Ba- cillus tuberigenus II, Gonnermann, Land- Avirtsch. Jahrb., 23, 1894, 656; Migula, Syst. d. Bakt., 2, 1900, 682.) From root nodules on a lupine. Bacillus gortynae Paillot. (Compt. rend. Acad. Sci., Paris, 157, 1913, 611.) From caterpillars of Gortyna ochracca. A coccobacillus. Bacillus gracilescens Henri ci. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 26.) From Swiss cheese. Bacillus gracilis Kern. CKern, Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1896. 421; not Bacillus gracilis Zimmermann, 658 MANUAL OF DETERMINATIVE BACTERIOLOGY Bakt. unserer Triiik- u. Nulzwasser, Chemnitz, 1, 1890, 50; Bacillus gracilior Migula, Syst. d. Bakt., 2, 1900, 664.) From the stomach and intestines of birds. Bacillus gracilis aerobiois \'aughan. (Amer. Jour. Med. Sei., 104, 1892, 187.) From water. Bacillus gracilis anaerobiescens \'aughan. (Amer. Jour. Med. Sci., 104, 1892, 187.) From water. Bacillus gracilis cudaveris Sternberg. (Sternberg, Man. of Bact., 1893, 733; Bacterium gracilis caiheris Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 84.) From the human liver. Bacillus grandis Trevisan. (BaciHe (ie Fair h, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 150; Trevi- san, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus gratnilaius Chester. {Bacil- lus aquaiilis solidvs Lustig, Diag. Bakt. om water. Bacillus salmoneus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 361; Bacterium salmoneus Chester, Ann. Rept . Del. Col. Agr. FAp. Sta., 9, 1897, 116.) From air. Bacillus salutarius Metchnikoff. (Aletfhnikoff, Maladies des hannetons (lu 1)1(', Odessa (in Russian), quoted from Paillot, L'infection chez les in- sectes, Paris, 1933, 123.) From diseased larvae of a beetle (Anisoplia auslriacn) . Bacillus sanguineus Schroeter. (In Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 156.) From stagnant water. Bacillus saponaceus Migula. (Bacil- lus lacHs saponacci Weigman and Zirn, Cent. f. Bakt., ]5, 1894, 464; Migula, Syst. d. Bakt., 2, 1900, 092.) From soapy milk. Bacillus saproiiciies Chester. {Bacil- lus saprogenes rini VI Kramer, Bak- teriol. Landwirtsch., 1890, 139; Chester, Man. Determ. l^act., 1901, 289.) From diseased wine. Bacillus saprogenes Trevisaii. (Bacil- lus saprogenes 1, Rosenbach, Mikro- organismen bei den Wundinfeclions- krankheiten des Menschen, Wiesbaden, 1884; Trevisan, I generi e le specie delle Batteriacee, 1889, 17; Bacterium sapro- genes Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 142.) From feces. Bacillus sarracenicolus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 357; Bacterium sarracenicolus (sic) Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 81 .) From a fresh leaf of the i)itcher ])lant (Sarraccnia purpurea). Bacillus scarldlinae Jamieson and Fa\- ingtoii. (Jamieson and Edington, Brit. Med. Jour., 1, 1887, 1265; Baeillus suda- minis Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) From the skin of scarlet fever patients. Bacillus scoticus Migula. (Bacillus der Grouse-disease, Klein, Cent. f. Bakt., 6, 1889, 36 and 593; ibid., 7, 1890, 81; Migula, Syst. d. Bakt., 2, 1900, 768; Bacillus telraonis Chester, Man. De- term. Bact., 1901, 221.) The cause of a disease of grouse (Lagopus scoticus) in England and Scotland. Bacillus secundus Trevisan. (Bacillus II, Leube, Arch. f. path. Anat., 100, 1885, 000; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From urine. Bacillus secmidus fullesi Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 359; Bacterium secundus fulessii (sic) Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 82; Bacterium seciindus ful- lesi Chester, ibid., 143.) From air. Dyai' regarded his organism as identical with Bacillus Xo. 2 of Fulles (Ztschr. f. Llyg., 10, 1891, 250) which was from soil. Bacillus seplicaemiac lophyri Shipero- vich. (Shiperovich, Protect. Plants I'kraine, 1925, 41-46; Abs. in Rev. Appl. Ent., A., 14, 1926, 209.) From larvae of sawflies (Di prion sertij'cr). Bacillus septicaemicus Trevisan. (Ba- cillus of septicemia of man, Klein, Micro- organisms and Disease, 1885, 84; Trevi- san, in DeToni and Trevisan, in Sac- cartlo, Sylloge Fungorum, 8, 1889, 945.) i-'rom blood and infected lymph glands. Bacillus septicus hominis Alironoff. (.Mironoff, Cent. f. Gynakol.. 1892, 42; Baclerium septicus hominis Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 143.) P'rom a case of septic infec- tion of the uterus. Regarded by Ches- ter (Man. Determ. Bact., 1901, 143) as a synonym of Pasteurella agrigena Tre- visan. Bacillus septicus vesicae Clado. (Bull, de la Soc. anatom. de Paris, 1887, 339.) From the urine of a person suffering from cystitis. Bacillus scriccus Zimmermann. (Bakt. unserer Trink- u. Xutzwjisser, Chemnitz, 2, 1894, 52.) From water. Bacillus .^crratus Migula. (Bacillus Xo. 14, Pansini, .\rch. f. path. .\nat., 122, 1890, 449; Migula, Syst. d. Bakt., 2, 1900, 658.) From sputum. Bacillus setosus Migula. (Bacillus Xo. XVIII, Adametz, Landwirtsch. Jahrb., FAMILY BACTBRIACEAE 669 18, 1889, 250; Migulu, Syst. d. Bakt., .?, 1900, 812.) From cheese. Bacillus silberschmidii Chester. (Ba- cillus der Fleischvergiftung, Silber- schmidt, Correspondenz-BUitt f. Scluvei- zer Aerzte, 1896, Xo. 8; Chester, Man. Determ. Bact., 1901, 212.) From poison- ous meat . Bacillus siniulatts Trevisan. (Bacille de Fair a, Babes, in Cornil and Babes, Les Bacteries, 2nd ed., 1886, 149; Trevi- san, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus singularis Losski. (Inaug. Diss., Dorpat, 1893, 45.) From garden soil. Bacillus siticulosiis Kern. (Arb. bakt . Inst. Karlsruhe, /, Heft 4, 1896. 423.) From the stomachs and intestines of birds. Bacillus sordidus Dj'ar. (Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 379; Bacterium sordidus Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 79.) Culture received by Dyar as Micrococcus sordidus from Krai's laboratory. Bacillus sordidus Kern. (.Vrb. Bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 396.) From the stomach of a bird. Bacillus sper7nophilinus Issatchenko. (Ein aus Zieselmjiusen ausgeschiedener Bacillus, Mereschkowsky, Cent, f . Bakt., 17, 1895, 742; Issatchenko, Scripta Bo- tanica Hort. Univ. Imp. Petropolitanae, Fasc. XV, 1897; quoted from ^Nligula, Syst. d. Bakt., .2, 1900, 618.) Apparently resembled Salmonelln tiiphimuriuDt . From ground squirrels {Spermophilus musicus). Bacillus spirans Weiss. (Arl). l)akt. Inst. Karlsruhe, 2, Heft 3. 1902, 222.) From bean infusions. Bacillus spwnosus Zimmermann. (Bakt. unserer Trink- u. Xutzwiisser, Chemnitz, 3, 1894, 28.) From water. Bacillus squamosus Pansini. (Arch, f. path. Anat., 122, 1890, 448.) From sputum. Bacillus strass)uan?ii Trevisan. {Ba- cillus albus cadaver is Strassmann and Strecker, Ztschr. f. Medicinalbeamte, 1888; Trevisan, I generi e le specie delle Batteriacee, 1889, 17; Bacterium albus cadaveris Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 102; Bacillus ca- daveris :Migula, Syst. d. Bakt., 2, 1900, 646.) From the blood of an infant. Bacillus striatus Doyen. (Bacillus urinae striatus Doyen, Jour. d. connaiss. medic, 1889, 107; Doyen, ibid., 108.) From urine. Bacillus striatus albus von Besser. (Ziegler's Beitrage, 4, 1889, 331.) Found in normal nasal mucus. Bacillus striatus flavus von Besser. (Ziegler's Beitrage, 4, 1889, 331; Bac- terium, striatus flavus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., .9, 1897, 111.) From nasal mucus. Rare. Bacillus stru77iitidis Migula. {Bacillus strumitis a, Tavel, Ueber die Aetiologie der Strumitis, Basel, 1892, 81; Migula, Syst. d. Bakt., 2, 1900, 741.) From a case of strumitis. Bacillus strumitis Tavel. (Tavel, 1889; see Viquerat, Ann. de ^licro- graphie, .2, 1889-1890,228.) From acute catarrhal strumitis. Bacillus subcoccoideus Migula. {Ba- cillus aquatilis sulcatus III, Weichsel- baum, Das osterreichische Sanitatswe- sen, 1889, Xo. 14-23; :\Iigula, Syst. d. Bakt., 2, 1900, 732.) From water. Bacillus subflavus Zimmermann. (Zimmermann, Bakt. unserer Trink. u. Xutzwiisser, Chemnitz, 1, 1890, 62; Bac- terium subflavus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 109.) From water. According to Chester (Man. Determ. Bact., 1901, 254), Bacillus flavescens Pohl is identical with this species. Bacillus subgastricus White. (U. S. Dept. Agr. Bur. Ent., Tech. Bui. 14, 1906, 23.) From intestinal contents of honey bee {Apis mellifera). While this does not appear to be the same as Bacil- lus gastricus Ford (see Steinhaus, Bac- teria Associated Extracellularly with Insects and Ticks. Minneapolis, 1942, 85), it may have been described by some 670 MANUAL OF DETERMINATIVE BACTERIOLOGY previous author as White does not indi- cate that he regards it as new. Bacillus subochraceus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 358; Bacterium subochraceus Chester, Ann. Rept. Del. Col. Ajr,-. Exp. Sta., 9, 1897, 110.) From air. Bacillus subrubiginosus Aligula. (Braunroter Bacillus, Maschek, Bakt. Untersuch. d. Leitmeritzer Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 836.) From water. Bacillus subsulcatus Migula. (Bacil- lus aquatilis sulcatus II, Weichselbaum, Das osterreichische SanitJitswesen, 1889, No. 14-23; Migula, Syst. d. Bakt., 2, 1900, 732.) From water. Bacillus sulcatus Chester. (Bacillus sulcatus liquefaciens Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 318; Bacterium sulcatus liquefaciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 97; Chester, Man. Determ. Bact., 1901, 2-13; not Bacillus sulcatus Migula, Syst. d. Bakt., ,?, 1900, 731.) From water. Bacillus sulcatus Migula. (Bacillus aquatilis sulcatus I, Weichselbaum, Das osterreichische Sanitatswesen, 1889, No. 14-23; Migula, Syst. d. Bakt., 2, 1900, 731.) From water. Bacillus sulfht/druqcuus Miquel. (Ann. de Micrographie, 1, 1888-1889, 369.) From sewage. Bacillus tardissimus DeToni and Tre- visan. (Bacillus fluidificans tardissimus Maggiora, Giorn. Soc. ital. d'Igiene, 11, 1889, 347; DeToni and Trevisan, in Sac- cardo, Sylloge Fungorum, 8, 1889, 967.) From the skin of the human foot. Bacillus tartricus Grimbert and Fi- quet. (Jour. Pharm. et de Chim., 6'" S^r., 7, 1898, 97; Compt. rend. Soc. Biol., 49, 1897, 962.) Decomposes d-tartrates. Probably identical with Aerobacier cloacae (Vaughn et al.. Jour. Bact., 52, 1946, 324). Bacillus telmatis Trevisan. (Bacillus saprogenes 2, Rosenbach, Mikroorganis- men bei den Wundinfectionskrankheiten des Menschen, Wiesbaden, 1884; Trevi- san, I generi e le specie delle Batteriacee, 1889, 14.) From perspiration of feet. Bacillus tenuis Doyen. (Bacillus urinae tenuis Doyen, Jour. d. connaiss. medic, 1889, 107; Doyen, ibid., 108; not Bacillus tenuis Migula, Syst. d. Bakt., 2, 1900, 587; Bacillus tenuatus Trevisan, in DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 948.) From urine. Bacillus tenuis apis White. (Jour. Path, and Bact., 24, 1921, 72.) From intestine of bee. Bacillus terrigenus Frank. (Berichte deutsch. botan. Gesellsch., 4, 1886, 000.) F'rom soil. Bacillus thermophilus Miquel. (Mi- quel, Ann. de Microgr., 1, 1888-1889, 6; not Bacillus thermophilus Chester, ]\Ian. Determ. Bact., 1901, 265; not Bacillus thermophilus Bergey et al.. Manual, 1st ed., 1923, 315.) From water, sewage, soil, etc. Bacillus thcta Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 375; Bacterium thcta C-hester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 144.) From air. Bacillus tingens Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 10.) From dead larvae of Orgyia pudibunda. Bacillus toluolicum Tausson. (Planta, 7, 1929, 735.) From soil. Oxidizes toluene. Bacillus toxigenus Chester. (Bacillus of ice cream poisoning, Vaughn and Perkins, Arch. f. Hyg., 27, 1896, 308; Chester, Man. Determ. Bact., 1901, 208.) From poisonous ice cream. Bacillus trambustii Kruse. (Tram- busti and Galeotti, Cent. f. Bakt., 11, 1892, 717; Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 319; Bac- terium trambusti Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 97.) From water. Bacillus tremaergasius Trevisan. (Ba- cille du mucus intestinal normal d, Babes, in Cornil and Babes, Les Bac- t(^ries, 2nd ed., 1886, 153; Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) From normal intestinal mucu.«. FAMILY BACTERIACEAE 671 Bacillus trimethylamin Beijeriiick. (Bot. Zeitung, 46, 1888, 726.) Bacillus truttae Mersch. (Quoted from Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 481.) Closely related to Bacterium salmonicida Leh- mann and Neumann. Bacillus tumidus Chester. (Anaerobe Bacillus I, Sanfelice, Ztschr. f. Hyg., U, 1S93, 268; Chester, INIan. Determ. Bact., ISOl, 265.) From putrefying flesh. Strict anaerobe. Bacillus uffrcduzzii Trevisan. (Bat- terio della setticemia salivare nei conigli , Bordoni-Uffreduzzi andDi-Mattei, Arch, per le scienze med., 10, 1886; Trevisan, see DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 951.) From normal human saliva. Bacillus ulna Cohn. (Beitr. z. Biol, d. Pflanz., 1, Heft 2. 1872, 177.) From water, air, etc. Bacillus umbilicatus Zimmermann. (Bakt. unserer Trink- u. Nutzwjisser, Chemnitz, :?, 1894, 6.) From water. Bacillus urinae iMigula. (Ein Harn- bacterium, Karplus, Arch. f. path. Anat.. 131, 1893, 211; Migula, Syst. d. Bakt., 2, 1900, 739.) From urine. Bacillus utpadeli Trevisan. (Bacillus aus Zwischendeckenfiillung, Utpadel, Arch. f. Hyg., 6, 1887, 359; Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) From the intestine. Bacillus vacuolalus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 357; Bac- teriuvi vacuolalus Chester, Ann. Rept. Del. Col. Agr. E.\p. Sta., 9, 1897, 81.) Frcm a trap of the carnivorous water plant, Vtricularia vulgaris. Bacillus vegetus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 399.) Frcm the stomach and intestines of birds. Bacillus velox Kern. (Arb. bakt . Inst . Karlsruhe, 1, Heft 4, 1896, 405.) From the stomach and intestines of birds. Bacillus venenosus brevis Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 192; Bacterium venenosus and Bacterium vene- nosus brevis Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 76.) From water. Bacillus venenosus Chester. {Bacillus venenosus invisibilis Vaughan, Amer. Jour. Med. Sci., 104, 1892, 192; Bacterium venenosus invisibilis Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 76; Chester, Man. Determ. Bact., 1901, 224; not Bacillus venenosus Vaughan, Amer. Jour. Med. Sci., 104, 1892, 191.) From water. Bacillus venenosus liquefaciens Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 193.) From water. Bacillus ventricosus Weiss. (Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 233; not Bacillus ve7itricosus Brede- mann and Heigener, Cent. f. Bakt., II Abt., 93, 1935, 102.) From vegetable infusions. Bacillus ventriculi Raczynski. (Diss, milit. medic. Acad. Petropolitanae Ru- teniae, 1888; abst. in Cent. f. Bakt., 6, 1889, 113.) From the stomach of a dog Bacillus vermiculosus Zimmermann (Zimmermann, Bakt. unserer Trink- u Nutzwasser, Chemnitz, 1, 1890, 40 Bacterium vermiculosus Chester, Ann Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 99.) From water. Bacillus versatilis DeToni and Trevi- san. (Bacillus A, Maggiora, Giorn. Soc. ital. d'Igiene, 11, 1889, 339; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 968.) From the skin of the human foot and from air. Bacillus vesiculiferus IMigula. {Bacil- lus strumitis 0, Tavel, Ueber die Aetiol- ogie der Strumitis, 1892, 110; IMigula, Syst. d. Bakt., 2, 1900, 741.) From a case of strumitis. Bacillus vialis Hansgirg. (Oesterr. bot. Zfschr., 1888, 6.) From roadside soil from near Prague. Bacillus viator Trevisan. (Bacille de Fair e. Babes, in Cornil and Babes, Les Bacteries, 2nded., 1886, 150; Trevisan, I generi e le specie delle Batteriacee, 1889, 20.) From air. Bacillus villosus Migula. {Bacillus aquatilis villosus Tataroff, Inaug. Diss., 672 MANUAL OF DETERMINATIVE BACTERIOLOGY Dorpat, 1891, 47; Migula, Syst. d. Bakt., 2, 1900, 828; not Bacillus villosas Keck, Inaug. Diss., Dorpat, 1890, 47.) From water. Bacillus vinicola Migula. {Bacillus saprogenes vini II, Kramer, Die Bak- terien in ihren Beziehungen zur Land- wirtschaft, 2, 1892, 136; Migula, Syst. d. Bakt., 2, 1900, 685.) From wine. Bacillus riniperda Migula. {Bacillus saprogenes vini I, Kramer, Die Bakterien in ihren Beziehungen zur Landwirt- schaft, 2, 1892, 135; Migula, Syst. d. Bakt., 2, 1900, 684.) From wine. Bacillus virens van Tieghem. (Bull. See. bot. France, 27, 1880, 175.) From aquatic plants. Bacillus viridans Zimmermann. (Bakt. unserer Trink." u. Xutzwasser, Chemnitz, 2, 1894, 22.) From water. Bacillus viridescens non-liquefaciens Ravenel. (Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 14; Bacterium viridescens non-liquefaciens Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 72 and 123.) From soil. Bacillus vulpinus von Iterson. (Cent . f. Bakt., II Abt., 12, 1904, 111.) From fresh garden soil, canal water. Bacillus wardii Chester. (Gas- and taint -producing bacillus in cheese curd, Moore and Ward, Cornell Univ. Agr. Expt. Sta., Bull. 158, 1899, 221-227; Chester, Man. Determ. Bact., 1901, 206.) From tainted, gassy cheese curd and from milk drawn directly from the udder. Presumably this was a colifoi'm or- ganism. Bacillus u'cckcri Trevisan. (Bacillus der Jequirity-Ophthalmie, de Wecker, 1882; see Flugge, Die Mikroorganismen, 2 Aufl., 1886, 279; Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From infusions of jequirity seed {Abri precaiorii) . Bacillus wesenbergii Chester. (Bacil- lus der Fleischvergiftung, Wesenberg, Ztschr. f. Hyg., 28, 1898, 484; Chester, Man. Determ. Bact., 1901, 247; not Bacillus wesenberg Castellani.) From meat which caused a meat poisoning outbreak. Closeh' related to Proteus vulgaris Hauser. Bacillus wichvianni Trevisan. (Gold- gelber Wasserbacillus, Adametz and Wichmann, Mitth. Oest. Vers. Stat. f. Brauerei u. Malz., 1, 1888, 49; Trevisan, I generi e le specie delle Batteriacee, 1889, 19; Bacillus chrrjseus Migula, Syst. d. Bakt., 2, 1900, 833.) From water. Bacillus zetaDyar. {T)ya.r, Ann. N. Y. Acad. Sci., 8, 1895, 369; Bacterium zeta Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 114.) From air. Bacillus zonatus Migula. (Bacillus No. 15, Pansini, Arch. f. path. Anat., 122, 1890, 450; Migula, Syst. d. Bakt., 2, 1900, 658.) From sputum. Bacillus zorkendorferi Migula. {Ba- cillus oogcnes Jnjdrosidfureus y, Zorken- dorfer. Arch. f. Hyg., 16, 1893, 385; Mi- gula, Syst. d. Bakt., ^, 1900, 696.) From hens' eggs. Bacillus zymoseus (Leube) Trevisan. {Coccobacillus zymogenes Leube, Arch. f. path. Anat., 1885; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From fermenting infusions. Bacterium acidi propionici Weigmann. (Weigmann, quoted from Pribram, Klas- sifikation der Schizomyceten, Leipzig und Wien, 1933, 76; Plocamobactcrium acidi propionici Pribram, idem.) Bacterium aeris ' Migula. {Bacillus aeris jniyiutissimus Kruse, in Fltigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 441; Bacterium aeris ininutissimus Chester, .Vnn. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 109; Migula, Syst. d. Bakt., 2, 1900, 445; Bacterium aeris-minutissimum Ches- ter, Man. Determ. Bact., 1901, 168.) From air. Bacterium aerogenes I and // Miller. (Miller, Deutsche med. Wochnschr., 12, 1886, 119; see Miller, Die Mikroorganis- men der Mundhohle, Leipzig, 1889, 261 and 262; not Bacterium aerogenes Ches- ter, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 53; Bacillus millerianus DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 952.) From the digestive tract of man. FAIVULY BACTERIACEAE 673 Bacterium agreste Lolinis. (Lohnis, Cent. f. Bakt., I Abt., Orig., 40, 1906, 177; Bacillus agrestis de Rossi, ^Microbiol, agraria e technica, Toriuo, 1927, 828; not Bacillus agrestis Werner, Cent. f. Bakt., II Abt., 87, 1933,468.) From soil. Bacterium agrigenum (Trevisan) Mi- gula. (Bacillus seplicus agrigenus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 257; Pasieurella agrigena Trevisan, I generi e le specie delle Batteriacee, 1889, 21; Bacterium septicus agrigenus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85; Migula, Syst. d. Bakt., 2, 1900, 372; Bacterium septicum Chester, Man. Determ. Bart.. 1901, 143.) From soil. Bacterium album Migula. (Weisser Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 35; :\Iigula, Syst. d. Bakt., 2. 1900, 419.) From water. Bacteriiim algeriensc ^ligula. (Gayoii and Dubourg, Ann. Inst. Past., 8, 1894, 108; Migula, Syst. d. Bakt., 2, 1900, 513.) Isolated in Algiers from wine where it causes a mannitol fermentation. Bacterium aliphaticum Tausz and Peter. (Cent. f. Bakt., II Abt.. ^5, 1919, 505.) From garden soil. Bacterium ali pJiaticum liqucfacicns Tausz and Peter. (Cent. f. Bakt., II Abt., 49, 1919, 505.) From garden soil. Bacterium allantoides (Klein) Chester. {Bacillus allantoides Klein, Cent. f. Bakt., 6, 1889, 383; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103.) Isolated as a culture contamination. Bacterium alutaceum Migula. (Gold- gelber chagrinierter Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 62; Migula, Syst. d. Bakt., 2. 1900, 464.) From water. Bacterium ambiguum Chester. (Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 11, 1900, 59; not Bacterium ambiguus Chester, ibid., 9, 1897, 71; not Bacterium amhigu\im Levine, Abst. Bact., 4, 1920, 15.) From soil. Bacterium amforeti Issatchenko. (Re- cherehes sur les microbes de I'Oc^an Glacial Arctique (in Russian), Petro- Srad, 1914, 237.) From sea water. Bacterium anaerobium JVIigula. (Fuchs, Inaug. Diss., Greifswald, 1890; Migula, Syst. d. Bakt., 2, 1900, 388.) Obligate anaerobe. Pyogenic. Pos- sibl}^ a spore-former. Bacterium auguillarum (Canestrini) Migula. (Bacillus anguillarum Canes- trini, Atti d. R. Instituto Veneto di Scienze, Ser. 7, 1892-93; Migula, Syst. d. Bakt., 2, 1900, 442.) From diseased eels in the valleys of Comacchio. Bacterium angustum Migula . (Lembke , Arch. f. Hyg., 26, 1896, 305; Migula, Syst. d. Bakt. ,2, 1900,474.) Fromfeces. Bacterium aphthosum (Kruse) Chester. (Bacillus der Mundsouche des ]\Ienschen, Siegel, Deutsch. med. Wochnschr., 1891, Xo. 49, 1328; Bacillus aphthosus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 427; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 85.) From the liver and kidneys of cattle affected with foot and mouth disease. Bacterium apis Xo. 1, X^o. 2 and X'o. 3, ^letalnikov and Kostritsky. (Compt. rend. Soc. Biol., Paris, 114, 1933, 1291.) From diseased bees (Apis mellifera.) Bacterium aquatile aurantiacum von Rigler. (Hyg. Rund., 12, 1902, 480.) From bottled mineral waters. Bacteriuvi aquatile citreum von Rigler. (Hyg. Rund., 12, 1902, 481.) From l)ottled mineral waters. Bacterium aquatile debile von Rigler. (Hyg. Rund., 12, 1902, 481.) From l)ottled mineral waters. Bacterium aquatile flavum von Rigler. (Hyg. Rund., 12, 1902, 480.) From bottled mineral waters. Bacterium aquatile luteum von Rigler. (Hyg. Rund., 12, 1902, 480.) From bottled mineral waters. Bacterium arborescens 7ion liquefaciens von Rigler. (Hyg. Rund., 12, 1902, 479; not Bacterium arborescens non-liquefa- ciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 103.) From bottled mineral waters. Bacterium arcticum Issatchenko . (Re- 674 MAXUAL OF DETERMINATIVE BACTERIOLOGY cherches sur Ics microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 148.) From sea water. Bacterium arthritidis Migula. (Schtil- ler, Berliner klin. Wochnschr., 1893, No. 36; Bacillus arthritidis chronicae Kruse, in Fltigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 287; Migiila, Syst. d. Bakt., 2, 1900, 443.) From a case of chronic arthritis. Bacterium asporiferum Migula. (Fltigge, Ztschr. f. Hyg., 17, 1894, 290; Anaerobier No. II, Kruse, in Fltigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 251; Migula, Syst. d. Bakt., 2, 1900, 446.) From milk. Bacterium aurantii (Viron) IVIigula. (Bacillus aurantii Viron, Compt. rend. Acad. Sci., Paris, 114, 1892, 179; Migula, Syst. d. Bakt., 2, 1900, 512.) Bacterium auraniium-roseum Honing. (Honing, Cent. f. Bakt., II Abt., 37, 1913, 373; Plocamobacterium aurantium Pribram, Klassifikation der Schizo- myceten, Leipzig und Wien, 1933, 77.) From fermenting tobacco. Bacterium aurescens (Franklaiid and Frankland) Migula. (Bacillus aurescens Frankland and Frankland, Philos. Trans. Royal Soc. of London, 178, 1887, B, 271; not Bacillus aurescens Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 8; Migula, Syst. d. Bakt., 2, 1900, 466.) From air. Bacterium aureum (Frankland and Frankland) Migula. (Baaillus aureus Frankland and Frankland, Philos. Trans. Royal Soc. of London, 178, 1887, B, 272; Migula, Syst. d. Bakt., 2, 1900, 480.) From air. Bacterium aureum (Adametz) Chester. (Bacillus aureus Adametz, quoted from Sternberg, Man. of Bact., 1893, 621; not Bacillus atireus Frankland and Frank- land, Philos. Trans. Roy. Soc. London, 178, 1887, B, 272; not Bacillus aureus Pansini, Arch. f. path. Anat., 122, 1890, 436; Bacillus aureo-flavus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 310; Bacterium atireo flams Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 109; Chester, ibid., 129; not Bac- terium aureum Migula, Syst. d. Bakt., 2, 1900, 480; Bacillus flavus Chester, Man. Determ. Bact., 1901, 255; not Bacillus flavus Fuhrmann, Cent. f. Bakt., II Abt., 1.9, 1907, 117; not Bacillus flavus Bergeyetal., Manual, Isted., 1923,286.) From water (Adametz); from the skin in cases of eczema (Tommasoli, Monats. f. prakt. Dermatol., 9.) Bacterium avium Chester. (Bacillus of roup in fowls, Moore, U. S. Dept. Agr. Bur. Animal Industry, Bull. 8, 1895; Chester, Man. Determ. Bact., 1901, 138.) From exudate of fowls in roup or diph- theria. Bacterium babesii Migula. (Bacillus septicus acuminatus Babes, Bakteriol- ogische Untersuchungen der septischen Prozesse des Kindesalters, Leipzig, 1889; see Eisenberg, Bakt. Diag., 3 Aufl., 1891, 327; Bacterium septicus acuminatus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 66; Migula, Syst. d. Bakt., 2, 1900, 507; Bacterium acuminatum Chester, Man. Determ. Bact., 1901, 119.) From blood and organs of a new-born infant with septicemia. Bacterium balbianii Billet. (Billet, Compt. rend. Acad. Sci., Paris, 107, 1888, 423; also in Bull. Sci. de la France et de la Belgique, 21, 1890, 108; Bacillus bal- bianii Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From sea water. Bacterium barentsianum Issatchenko. (Recherches sur les microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 155.) From sea water. Bacterium beijerincki Issatchenko. (Recherches sur les microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 157.) From sea water. Bacterium benzoli a and b Wagner. (Ztschr. f. Garungphysiol., 4, 1914, 289.) From soil. Utilize benzene and certain benzene derivatives. Bacterium besseri Migula. (Besser, Cent. f. Bakt., 13, 1893, 590; Migula, Syst. d. Bakt., 2, 1900, 508.) From smallpox. Bacterium betae viscosum Panek. (Bull. Acad. Sci. Cracovie, ;, 1905, 5.) FAMILY BACTERIACEAE 675 From fermenting beets. Reported to liquefy agar-gelatin (Biernacki, Cent. f. Bakt., 29, 1911, 166). Stanier (Jour. Bact., 42, 1941, 548) thinks this was a heterofermentative Lactohacillus. Bacterium bossonis Chester. (Bacillus fiber eine neue Infektionskrank. des Rindviehs, Bosso, Cent. f. Bakt., 22, 1897, 537 and 23, 1898, 318; Chester, Man. Determ. Bact., 1901, 153.) Associated with an infectious disease of cattle. Bacterium boutro^ixii (Trevisan) De- Toni and Trevisan. (Micrococcus ca- pable d'acetifier I'alcohol, Boutroux, Ann. Inst. Past., 2, 1888, 209; Bacillus boutrouxii Trevisan, I generi e le specie delle Batteriacee, 1889, 16; DeToni and Trevisan, in Saccardo, S3'lloge Fung- orum, 8, 1889, 1021.) From alcoholic infusions. Bacterium bovis iNIigula. (Pncumo- bacillus liquefaciens bovis Arloing, Compt. rend. Acad. Sci., Paris, 99, 18—, 109 and 116; Bacillus pneumomcus liquefaciens Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 288; Bacterium prieu- monicus liquefaciens Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 99; Migula, Syst. d. Bakt., 2, 1900, 442; Bacterium pneumonicum Chester, ]\Ian. Determ. Bact., 1901, 158.) From the exudate of lung plague in cattle. A Gram-positive coccus-like bacterium. Bacterium brassicae Conrad. {Bac- terium brassicae acidae Lehmann and Conrad, in Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 232; Con- rad, Arch. f. Hyg., 29, 1897, 82; not Bacterium brassicae Wehmer, Cent. f. Bakt., II Abt., 10, 1903, 628; not Bac- terium brassicae Migula, Syst. d. Bakt., 2, 1900, 296 {Bacillus brassicae Pommer, Mitt, botan. Inst. Graz, 1, 1886, 95); Bacillus brassicae Migula, loc. cit., 737.) From sauerkraut. Bacterium breitfussi Issatchenko. (Recherches sur les microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 152.) From sea water. Bacterium brevissimum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 227.) From vegetable infusions. Bacterium brunneofiavum (Dyar) Ches- ter. {Bacillus brunneoflavus Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 362; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 112.) Culture received from Krai's laboratory as Micrococcus brunneus. Bacterium bullosum Migula. (Bacillus No. 18, Pansini, Arch. f. pathol. Anat. u. Physiol., 122, 1890, 451; Migula, Syst. d. Bakt., 2, 1900, 415.) From feces. Bacteriuin cadaveris (Sternberg) Ches- ter. {Bacillus cadaveris Sternberg, Man. of Bact., 1893, 492; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 126.) From liver and kidneys of yellow fever cadavers. Anaerobe. Bacterium canalis Migula. (Kapsel- tragender Kanalbacillus, Mori, Ztschr. f. Hj^g., 4, 1888, 52; Bacillus canalis cap- sulatus Sternberg, Man. of Bact., 1893, 476; Bacterium canalis capsulatus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta. , 9, 1897, 130; Migula, Syst. d. Bakt., 2, 1900, 351.) From sewage. Bacterium canalis parvum (Sternberg) Chester. {Bacillus canalis parims Stern- berg, Man. of Bact., 1893, 476; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 130.) Obtained by Mori (1888) from sew^age. Bacterium carneum (Kruse) Chester. (Fleischfarbiger Bacillus, Tils, Ztschr. f. Hyg., 9, 1890, 294; Bacillus carnicolor Frankland and Frankland, Microor- ganisms of Water, 1894, 477; Bacillus carneus Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 304; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 113.) From water. Bacterium carnosum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 448.) From the intestines of birds. Bacterium cartilagineum (Olsen-Sopp) Buchanan and Hammer. •(Bacillus car- tilagineus Olsen-Sopp, Cent. f. Bakt., II Abt., 33, 1912, 49; Buchanan and Hammer, Iowa Sta. Agr. Exp. Sta., Res. Bull. 22, 1915, 271.) From slimy or ropy sour milk called false taette. 670 MAXUAL OF determixativp: bacteriology Bacteriuni caseicola Aligula. (Bacil- lus No. XII, Adametz, Landw. Jahrb., 18, 1889, 245; Migula, Syst. d. Bakt., 2, 1900, 475.) From cheese. Bacterium castellum Henrici. (Arh. bakt. Inst. Karlsruhe, 1, Heft 1, 1S94, 38.) From cheese. Bacteriiivi cntenula Dujardin. (Du- jardin. Hist, natur. des zooph., 1841; Bacillus catenula. Trevisan, I generi e le specie delle Batteriacee, 1889, 18; not Bacillus catenula Migula, Syst. d. Bakt., 2, 1900, 588.) Fr.om rice paddies and swamps. Bacterium caratum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 449.) From the intestines of birds. Bacterium cavernae Aligula. {Bacillus caveruae minutissijuus Kruse, in Fliigge, Die Mikroorganismen, 3 Auti., 2, 1896, 440; Pfciffer und Beck, Deutsch. med. Wochnschr., 1892, No. 21; Aligula, Syst. d. Bakt., 2, 1900, 509.) From human tuberculosis. Bacterium caviaejorluitum (Sternberg) Chester. {Bacillus caviae furtuitus Sternberg, Alan, of Bact., 1893, 650; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74.) From the liver of a 3'ellow fever cadaver. Bacterium cavicida havaniensis (>Stern- berg) Chester. {Bacillus cavicida hava- niensis Sternberg, Alan, of Bact., 1893, 425; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74.) From, the in- testine of a yellow fever cadavei-. Bacterium centricum Aligula. (Huber, Armin, Arch, f . pathol. Anat. u. Physiol., 134, 1893, 216; Aligula, Syst. d. Bakt., 2, 1900, 390; not Bacterium concentricum , a typographical error, see Aligula, ibid., page V.) From a case of cystitis. Bacterium cerinum Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 50.) From cheese. Bacterium chlorinum Engelmann. (Bot. Zcitung, 1882, 324.) Green pig- ment. Bacterium cholocjencs (Kruse) Chester. (Colonbacillus, Stern, Deutsche med. Wochnschr., 1893, 613; Bacillus cholo- (jcncs Kruse, in Fliigge, Die Alikroor- ganismen, 3 Aufl., 2, 1896, 374; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 69.) From a case of purulent meningitis. Bacterium chryseum Aligula. (Bacil- lus nova species II, Freund, Inaug. Diss., Erlangen, 1893, 37; Aligula, Syst. d. Bakt., 2, 1900, 477.) Chromogenic bacterium from tlie mouth cavity. Bacterium chrysogloea Zopf. (Zopf, in Overbeck, Nova Acta d. kais. Leop.- Carol. Akad. d. Naturf., 55, 1891, No. 7; Bacillus chrysogloia (sic) Zimmermann, Bakt. unserer Trink- u. Nutzwasser, 2, 1894, 12.) From water. Bacterium citrcnui (Frankland and Frankland) AligiUa. {Bacillus citreus Frankland and Frankland , Philos. Trans . Royal Soc. of London, 178, 1887, B, 272; Aligula, Syst. d. Bakt., 2, 1900, 459.) From air. Bacterium culi apinm Serbinow. (Jour. Alicrobiol. Petrograd., 2, 1915, 19.) From honey bees {Apis mellifera) . Bacterium coli sittiilis (Sternberg) Chester. {Bacillus coli similis Stern- berg, Alan, of Bact., 1893, 650; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 132.) From a human liver. Bacterium culloideum Aligula. {Bac- terium butyri colloideum Lafar, Arch. f. Hyg., 13, 1891, 17; Aligula, Syst. d. Bakt., 2, 1900, 409.) From butter. Bacterium comes Berstcyn. (Arb. l)akt. Inst. Karlsruhe, 3, 1903, 93.) From soil . Bacteriuui cotnpaclum (Kruse) Aligula. {Bacillus cotupactiis Kruse, in Fliigge, Die Alikroorganismen, 3 Aufl., 2, 1896, 353; Aligula, Syst. d. Bakt., 2, 1900, 438.) From air. Bacterium concentricum Kern. (Arb. l)akt. Inst. Karlsruhe, 1, Heft 4, 1896, 437.) From the intestines of birds. Bacterium conjunctivitidis (Kruse) Ali- gula. (Koch, Berichte aus Aegypten an den preuss. Staatsminister des In- nern; see Arb. a. d. kaiserl. Gesund- heitsamte, 3, 1887; Kartulis, Cent. f. Bakt., 1, 1887, 289; Bacillus aegyptius FAlVnLY BACTERIACEAE 677 Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Bacillus conjuncti- vitidis Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 440; Bac- terium conjunctivitis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 67; Mi- gula, Syst. d. Bakt., 2, 1900, 509; not Bacterium conjunctivitidis Chester, ]Man. Determ. Bad., 1901, 120; Bacterium aegyptiu/n Chester, ibid., 121.) Asso- ciated with conjunctival catarrh in Egypt . Bacterium corticate (Haenlein) ^Sligula. {Bacillus corticalis Haenlein, Deutsch. Gerberzeitung, 1894, Xo. 18-34; :\IiguUi, Syst. d. Bakt., 2, 1900, 449.) Found on pine bark; in acid dyeing-liquor. Bacterium crenatum Weiss. (Arlj. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 221.) From fermenting malt. Bacterium cristalliferum Gicklhorn. (Cent. f. Bakt., II Abt.. 50, 1920, 420.) A sulfur bacterium from soil. See Manual, 5th ed., 1939, 86 for a descrip- tion of this organism. Bacterium cuticularis (Tils) Chester. (Bacillus cuticularis Tils, Ztschr. f. Hyg., 9, 1890, 293; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 105.) P'rom water. Bacterium debile Berstj'n. (Arb. bakt . Inst. Karlsruhe, 3, 1903, 96.) From soil. Bacterium delendae-muscae Roubaud and Descazeaux. (Compt. rend. Acad. Sci., Paris, 177, 1923, 716.) From fly larvae (Slomoxys calcitrans and Musca domestica) . Bacterium deliciise Honing. (Cent, f. Bakt., II Abt., 37, 1913, 377.) From tobacco plants in Sumatra. Bacterium diatrypeticum ]\Iigula . {Bacillus diatrypeticiis casei Baumann, Landwirtsch. Versuchsstationen, ^2, 1893, 181; Migula, Syst. d. Bakt., 2, 1900, 404.) From cheese. Bacterium enclielys Ehrenberg. (Ehrenberg, Abhandl. d. Akad. d. Wissensch. zu Berlin, 1830, 61; Bacillus enchelys Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) From water. Bacterium endocarditidis Migula. {Bacillus endocarditidis capsulatus Weichselbaum, Beitr. z. pathol. Anat. u. z. allgem. Pathol., 4, 1887, 197; Migula, Syst. d. Bakt., .?, 1900, 359.) Found in the aorta, the left ventricle, the spleen and kidneys of cadavers. Bacterium endometritidis (Kruse) Chester. {Bacillus endometritidis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 432; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 88.) From a liver abscess. Bacterium, endometritis canis Meyer. (Meyer, quoted from Pribram, Klassi- fikation der Schizomyceten, Leipzig und Wien, 1933, 77; Plocamubacterium endometritis Pribram, idem.) From a case of endometritis in a dog. Bacterium enterocoliticum Schleifstein and Coleman. (A motile. Gram-nega- tive bacillus, Schleifstein and Coleman, X. Y. State Jour. Med., 39, 1939, 1749; Ann. Rept. Div. Lab. and Res., X. Y. State Dept. Health for 1943, 56.) From lesions about the face, from an ulcer in the intestine and from the intestinal contents. Resembles Bacillus lignieri and Pasteurella pseudotuberculosis. Bacterium erythromyxa (Zopf) IMigula. {Micrococcus {Staphylococcus) erythro- myxa Zopf, Ber. d. deutschen bot. Gesellsch., 1891, 22; Rhodococcus ery- thromyxa Zopf, loc. cit.; Migula, Syst. d. Bakt., 2, 1900, 487; Bacillus erythro- myxa Matzuschita, Bakt. Diag., 1902, 389.) Frequently listed as a Micro- coccus. Bacterium exantheniaticum (Kruse) Chester. (Bacille, Babes and Oprescu, Ann. Inst. Past., 5, 1891, 273; Bacillus exanthematicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 426; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 87.) From a case of hemor- rhagic septicemia. Bacterium fausseki Issatchenko. (Re- cherches sur les microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 157.) From sea water. Bacterium ferophilum (sic) Migula. 678 MANUAL OF DETERMINATIVE BACTERIOLOGY (Die ferrophilen Bakterien, Marpmann, Cent. f. Bakt., II Abt., 4, 1898, 21; Migula, Syst. d. Bakt., 2, 1900, 455 and 1058.) Isolated during studies on black discoloration of cheese. Bacterium finitimum Chester. {Ba- cillus finitimus ruber Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 361; Bacterium finiti- mus ruber Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 116; Chester, Man. Determ. Bact., 1901, 177.) From air. Bakterium flaveitm Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 15.) From meat extract. Bacterium flavocoriaceum (Eisenberg) Chester. (Schwefelgelber Bacillus, Adametz and Wichmann, Die Bakterien der Trink- und Nutzwiisser, Mitt. Oest. Versuchsstat. f. Brauerei u. Malz., Wien, Heft 1, 1888, 49; Bacillus flavo- coriaceus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 144; Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 112.) From water. Bacterium flaTofuscum Migula. (Xo. 9, Lembke, Arch. f. Hyg., 26, 1896, 304; Migula, Syst. d. Bakt., 2, 1900, 479.) From meat. Bacterium flavum Issatchenko. (Re- cherches sur les microbes de 1 'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 151.) From sea water. Bacterium foliicola (Miehe) de Jongh. (Bacillus folii cola Miehe, see Jahrb. wiss. Bot., 53, 1914, 1; ibid., 58, 1919, 29; de Jongh, On the Symbiosis of Ardisia crispa. Thesis, Univ. Leiden, 1938, 33.) A bacterial symbiont isolated from germinating seeds and embryos. Bacteriujn freundii Migula. (Bacillus nova species I, Freund, Inaug. Diss., Erlangen, 1893, 31; Migula, Syst. d. Bakt., 2, 1900, 472.) From the mouth cavity. Bacterium fungoides (Tschistowitsch) Migula. (Bacillus fungoides Tschisto- witsch, Berl. klin. Wochnschr., 1892, 513; Migula, Syst. d. Bakt., 2, 1900, 391.) From pus. Bacterium fuscum (Fliiggc) ]Migula. (Bacillus fuscus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 290; Migula, Syst. d. Bakt., 2, 1900, 463.) From water. Bacterium gamma (Dyar) Chester. (Bacillus gamma Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 367; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 106.) From air. Bacterium gammari Vejdovsky. (Cent. f. Bakt., II Abt., /;, 1904, 484.) From sections of a fresh water crustacean (Gammarus zschokkei). Cells exhibit nuclei, showing mitosis. Bacterium gelechiae No. 1 and No. 2, Metalnikov and Metalnikov. (Compt. rend. Acad. Agr., France, 18, 1932, 204.) From dead and dying larvae of a moth (Gelechia gossypiella) . Bacterium gelechiae No. 5, ^letalni- kov and Meng. (Compt. rend. Soc. Biol., Paris, 113, 1933, 170.) From dead larvae of a moth (Gelechia gossypiella) . Bacterium gemmijorme Migula. (Lembke, Arch. f. Hyg., 29, 1897, 313; Migula, Syst. d. Bakt., 2, 1900, 391.) From intestinal contents. Bacterium gibbosum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 230.) From fermenting beets and malt. Bacterium gingivae pyogenes Miller. (Miller, Die Mikroorganismen der Mund- hohle, Leipzig, 1888, 217; Bacillus gingi- vae pyogenes Sternberg, Manual of Bact., 1893, 471.) From an alveolar abscess. Bacterium gingivitidis (Kruse) Migula. (Babes, Deutsch. med. Wochnschr., 1893, 1035; Bacillus gingivitidis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl. , 2, 1896, 427; Migula, Syst. d. Bakt., ^, 1900, 393.) Isolated in an epidemic of scurvy in Jassy. Bacterium gliscrogcnum Malerba and Sanna-Salaris. (Malerba and Sanna- Salaris, Lavori eseguiti nell'Istituto fisiol. di Napoli, 2, 1888, 13 and 95; Bacil- lus gliscrogenus Trevisan, I generi e le specie delle Batteriacee, 1889, 14.) From urine. Bacterium gonnermanni Migula. (Ba- cillus tuberigenus 6 Gonnermann, Land- FAMILY BACTERIACEAE 679 wirtsch. Jahrb., 23, 1894, 657; Aligula, Syst. d. Bakt., 2, 1900, 418.) From root ncdules of lupine. Bacterium gracilescens Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 259.) From fermenting asparagus and malt . Bacterium gracillimum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 235.) From bean and asparagus in- fusions. Bacterium granulatum Henrici. (Hen- rici, Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 33; not Bacterium granulatum Chester, Man. Determ. Bact., 1901, 189.) From cheese. Bacterium granulosum Weiss. (Weiss, Arb. bakt. Inst. Karlsruhe, 5, Heft 3, 1902, 212; not Bacterium granulosum Lehmann and Neumann, Bakt. Diag., 5 Aufl., 2, 1912, 306.) From vegetable infusions. Bacterium gryllotalpae ^letalnikov and Meng. (Compt. rend. Acad. Sci., Paris, 201, 1935, 367.) From diseased larvae of the cricket (Gryllotalpa gryllotalpa) . Bacterium gummosum Ritsert. (Rit- sert, Ber. d. pharmaz. Gesell., /, 1891, 389; abst. in Cent. f. Bakt., 11, 1892, 730; Bacillus gumynosus IMigula, Sj'st. d. Bakt., 2, 1900, 873.) A mi.xture of a spore-forming rod and a streptococcus. See Bacillus gummosus Happ and Micro- coccus gu77i7nosus Happ. Bacterium halans (Zimmermann) ^li- gula. (Bacillus halans Zimmermann, Die Bakterien unserer Trink- u. A'utz- wasser, Chemnitz, 2, 1894, 54; Migula, Syst. d. Bakt., 2, 1900, 429.) From water. Bacterium hehetisiccus Steinhaus. (Jour. Bact., ^2, 1941, 762 and 773.) From the walking stick {Diapheromera femorata) . Bacterium herbicola a aureum Gei- lingcr. (Mitteil. a. d. Gebiete d. Lebensmitteluntersuchungen u. Hyg., 12, 1921, 262.) From corn meal. This is a variety of Bacillus herbicola Burri and Diiggeli. Bacterium herbicola rubrum Diiggeli. (Duggeli, Cent. f. Bakt., II Abt., 12, 1904, 605; Bacterium herbicola /3 rubrum, Lehmann and Neumann, Bakt. Diag., 4 -\ufl., 2, 1907, 356.) From germinating plants, roots and barley seeds. Bacterium hexacarbovorum Stormer. (Jahresber. d. Vereinigg. f. angew. Botanik, 5, 1907, 116.) From soil. Utilizes benzene and certain benzene derivatives. Bacterium hidium Goobin. (Russian Health Resort Service, 5, 1923, 3.) At- tacks ethane and other hydrocarbons. Bacterium hirudinicolicum Lehmen- sick. (Cent. f. Bakt., I Abt., Orig., 1^7, 1941, 317; see Biol. Abst., 18, 1944, No. 6761.) Symbiotic in the intestines of Hirudo officinalis and H. medicinalis. Bacterium hoshigaki var. glucuronicum II and III Takahashi and Asai. (Cent, f. Bakt., II Abt., S7, 1933, 395 and 405.) From dried persimmons (hoshigaki). Bacterium infecundum Chester. (Ba- cillus filiformis havaniensis Sternberg, Man. of Bact., 1893, 650; Bacterium fili- formis havaniensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 126; Chester, Man. Determ. Bact., 1901, 184.) From the liver of a yellow fever cadaver. Anaerobic. Bacterium inocuum Chester. (Wilde, Wien. klin. Wochnschr., 1892, No. 1-2; Bacillus lactis inocuus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 352; Bacterium lactis inocuus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 82; Chester, Man. Determ. Bact., 1901, 138.) From milk. Bacterium intrinsectum Steinhaus. (Jour. Bact., ^2, 1941, 764 and 774.) From an unidentified leaf beetle. Bacterium iogenum Baumgartner. (Deutsche Monatschr. f. Zahnheilk., 27 , 764; Bacterium jogenum Baumgartner, Ergeb. d. gesam. Zahnheilk., 1, 1911, 752 and 779; B. iogenum Kligler, Jour. Allied Dent. Soc, 10, 1915, 152.) From the mouth. Regarded as identical with Jodococcus vaginatus Miller by Kligler (loc. cit.). Bacterium keratomalaciae Migula. (Bacillus septicus keratomalaciae Babes, 680 MANUAL OF DETERMINATIVE BACTERIOLOGY Bakteriol. Untersuch. d. sept. Prozesso d. Kindesalters, Leipzig, 1889; INIigula, Syst. d. Bakt., £, 1900, 363.) From an infected cornea. Bacterium knipowitchi Issatchenko. (Recherches sur les microbes de 1 'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 150.) From sea water. A phosphorescent bacterium found to be pathogenic for the mealworm {Tenebrio molitor) by Pfeiffer and Stammer (Ztschr. f. Morph. u. Okol. d. Tierc, 20, 1930, 157). Bacterium kralii (Dyar) Chester. {Bacillus kralii Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 376; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 93; not Bacterium kralii Chester, Man. Determ. Bact., 1901, 166.) Received as Bacillus butyricus from Krai's laboratory by Dyar. The 1900 Krai Catalogue lists cultures of Bacillus butyricus Botkin and Bacillus butyricus Hueppe. As Dyar found that the characters of his culture differed from those of Bacillus butyricus Hueppe, Dyar's culture was probably Bacillus butyricus Botkin. Bacterium kralii Chester. (Bacillus fuscus liquefaciens Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 375; Bacterium fuscus liquefaciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 108; Chester, Man. Determ. Bact., 1901, 166.) Re- ceived as Bacillus fuscus from Krai's laboratory by Dyar who also found it in air. The 1920 Krai catalogue lists Bacil- lus fuscus Fliigge syn. Bacterium brun- neum Schrotter (sic); braunrother Ba- cillus Maschek. Bacterium laerii Migula. (Bacillus viscosus No. 1, van Laer, Extrait des m^moires couronn^s et autres mdmoires, Acad. Royale de Belgique, 1889, 36; see Kramer, Bakteriol. in ihren Beziehungen zur Landwirtsch., 2, 1892, 119; Bacillus viscosus cerevisiae Kruso, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 359; Bacterium viscosus cerevisiae Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 78; Migula, Syst. d. Bakt., S, 1900, 402; Bacterium viscosum Chester, Man. Determ. Bact., 1901, 128.) From beer, yeast, air, bread. Causes a slimy fer- mentation . Bacterium laevolacticum Migula. (Ba- cillus acidi laerolactici Schardinger, Monatsh. f. Chemie, 11, 1890, 544; Mi- gula, Syst. d. Bakt., 2, 1900, 406; Bac- terium acidi laevolactici Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 178.) From milk. Bacterium laminariae Billet . (Compt . rend. Acad. Sci., Paris, 106, 1888, 293; Billetia laminariae Trevisan, I generi e le specie delle Batteriacee, 1889, 11; Kurthia laminariae De Toni and Trevi- san, in Saccardo, Sylloge Fungorum, 8, 1889, 931.) From rotting sea weed. The type species of the genus Billetia Trevisan. Bacterium langkatensc Honing. (Cent. f. Bakt., II Abt., 37, 1913, 381.) From tobacco plants in Sumatra. Bacterium largum (v. Klecki) Migula. (Bacillus largus v. Klecki, Ann. Inst. Past., 9, 1898, 728; Migula, Syst. d. Bakt., ^, 1900, 448.) From the intestines of dogs. Bacterium lepierrei Chester. (Bacille fluorescent pathogene, Lepierre, Ann. Inst. Past., 9, 1895, 643; Chester, Man. Determ. Bact ., 1901, 182.) From cistern water. Bacterium lelhalis (Babes) Chester. (Proteus lethalis Babes, Progres Medical Roumain, 1889; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 5, 1897, 104; Bacillus lethalis Chester, ]\Ian. Determ. Bact., 1901, 249.) From lung gangrene in man. Bacterium leucaemiae Migula . (Lucet , Jahresber. li. Fortschr. in d. Lehre v. d. path. Mikroorg., 7, 1891, 319; Bacillus leucaemiae canis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 285; Bacterium leucaemiae canis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 119; Migula, Syst. d. Bakt., ^, 1900, 442; Bacillus leucaemiae Chester, Man. Determ. Bact., 1901, 264.) From a dog with leukemia. Bacterium linibatuin Migula. (Bac- terium limbatum acidi lactici Marpmann, FAMILY BACTERIACEAE 081 Ergaiizunfishoflo d. Centralb. t'. allg. (Jesuidhrilspflege, ^, 122; Bacillus lim- hatitti iicidi lactici Sternberg, Man. of Bact., 1S93, 645; Migula; Syst. d. Bakt., 2, 1900, 407.) From fresh milk. Bacterium lineola (Miiller) C'ohu. {Vibrio lineola ]\Iuller, Vermium Ilis- icria, 1773, 39; Cohn, Beitr. z. Biol. d. Pfianz., /, Heft 2, 1872, 170; Bacillus lineola Trevisan, I generi e le specie delle Batteriaree, 1889, 18.) From stagnant water, infusions, etc. Bacterium lint Migula. (Winogiad- sky, Compt . rend. Acad. Sci., Paris, 121 , 1893, 742; Migula, Syst. d. Bakt., .?, 1900, 513.) From retting hemp. Bacterium iinkoi Issatchenko. (lle- cherches sur les microbes de I'Occan Glacial Arctique (in Russian). Petro- grad, 1914, 154.) From sea water. Bacterium liquefaciens communis (Sternberg) Chester. {Bacillus lique- faciens communis Sternberg, ]\Ian. of Bact., 1893, 686; Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 91; Bacillus communis Migula, Syst. d. Bakt.,,^, 1900, 725; not Bacillus communis Jackson, Jour. Inf. Dis., 8, 1911, 241.) From the feces of yellow fever patients. Bacterium litoreum Warming. (Warm- ing, Danmarks Kyster levende Bak- terier, 1875, 39S; Bacillus litoreus Trevi- san, I generi e le specie delle Batteriacee, 1889, 18.) From sea water. Bacterium loculosum Migula. (Fach- erbacillus, Clauss, Inaug. Diss., Wiirz- burg, 1889, 27; Migula, Syst. d. Bakt., 2, 1900, 408.) From milk. Bacterium luceti ^ligula. (Lucet, Ann. Inst. Past., 3, 1889, 401; Bacillus cuniculicida thermophilus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 418; Migula, Syst. d. Bakt., 2, 1900, 507; Bacterium cuniculicida thermo- philus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 83; Bacterium cunicidicida Chester, Man. Determ. Bact., 1901, 140.) Associated with an epizootic in rabbits and guinea pigs. Bacterium ludwigi Karlinski. (Hyg. liundschau, 5, 1895, 685.) From the water of the hot springs at Ilidze in Bosnia. Bacterium luteolum Henrici. (Henrici, .\rb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 51; Migula, Syst. d. Bakt., 2, 1900, 455.) From cheese. Bacterium, lutescens ^Migula. (Dcr gelbe Bacillus, Lustig, Diag. d. Bakt. d. Wassers, 1893, 78; Migula, Syst. d. Bakt., 2, 1900, 476.) From water. Bacterium luieum Adametz. (List, Inaug. Diss., Leipzig, 1885, 53; Adametz, Bakt. Nutz- u. Trinkwasser, Mitteil. d. ostcrr. Versuchsstation fiir Brauerei und Malzerei in Wien, 1888, 48.) From the stomach contents of sheep and from water. Bacterium margurineum ^ligula. (iNIargarinbacillus a, Jolles and Winkler, Ztschr. f. Hyg., 20, 1895, 102; Migula, Syst. d. Bakt., 2, 1900, 410.) From margarine. Bacterium marinum Issatchenko. (Recherches sur les microbes de 1 'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 238.) From sea water. Bacterium maydis Maiocchi. (Mai- occhi, Bollet. d. Accad. medic, d. Roma, October, 1881 ; Bacillus maydis Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From corn (maize) infusions. Bacterium medanense Honing. (Cent, f. Bakt., II Abt., 37, 1913, 382.) From the peanut plant {Arackis hypogaea). Bacterium melolonthae liquefaciens Paillot . (Compt . rend . Soc . Biol . , Paris , 68, 1916, 1102.) From the cockchafer {Mclolontha melolontha). According to the author's system of nomenclature, this is presumably a synonym of Bacillus melolonthae liquefaciens a. Bacterium meningitidis (Neumann and Schaffer) Chester. {Bacillus meningi- tidis purulentae Neumann and SchaefTer, Arch. f. path. Anat., 109-, 1887, 477; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 71; Bacillus neximanni Mi- gula, Syst. d. Bakt., 2, 1900, 751; not Bacillus neumanni Herter, in Just, Botan. Jahresber., 2 Abt., 39, Heft 4, 1915, 748; Bacillus meningitidis Chester, 682 MANUAL OF DETERMINATIVE BACTERIOLOGY Man. Determ. Bact., 1901, 213.) From pus from an individual who died of puru- lent meningitis. Bacterium merismopedioides Zopf. (Zopf, Die Spaltpilze, 1 Aufi., 1883, 56; Bacillus synchyseus Trevisan, I generi e le specie delle Batteriacee, 1889, 18; Bacterium synchyseus DeToni and Trevi- san, in Saccardo, Sylloge Fungorum, 8, 1889, 1022.) From canal water. Bacterium microsporum Trevisan. (Trevisan, Rendic. d. Instit. Lombardo. Ser. 2, 13, 1879; Bacilhis microtis Trevi- san, I generi e le specie delle Batteriacee, 1889, 18; Bacterium microtis DeToni and Trevisan, in Saccardo, Sylloge Fungo- rum, 8, 1889, 1025.) From water and putrefying infusions. Bacterium minutmn (Zinnnermann) Migula. (Bacillus minutus Zimmer- mann, Die Bakterien unserer Trink- und Nutzwasser, Chemnitz, 2, 1894, 56; Mi- gula, Syst. d. Bakt., ;3, 1900, 423.) From water. Bacterium monachac von Tubeuf. (v. Tubeuf, Forstlicli - naturwissensch. Ztschr., 1, 1892, 34; Bacillus monachae Migula, Syst. d. Bakt., 2, 1900, 742.) From the larvae of a moth (Lymantria monacha) . Bacterium multipcdiculum (Jliigge) Chester. (Bacillus multipediculus Flligge, Die Mikroorganismen, 2 Aufl., 1886, 323; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 104.) Isolated frequently as a contamination on potato media. Bacterium muripcstifer (Kruse) Ches- ter. (Bacillus dcr Mauseseuche, Laser, Cent. f. Bakt., 11, 1892, 184; Bacillus muripcstifer Kruse, in Flligge, Die Mikroorganismen, 3 Aufl., 2, 1896, 432; Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 87.) From the spleen of a field mouse. Associated with a plague of field mice. Bacterium nacreaceum (Zimmermann) Migula. (Perlmutterglanzender Bacil- lus, Keck, Inaug. Diss., Dorpat, 1890, 40; Eberbach, Inaug. Diss., Dorpat, 1890; Bacillus nacreaceus Zimmermann, Die Bakterien unserer Trink- und Nutz- wasser, Chemnitz, 2, 1894, 34; Migula, Syst. d. Bakt., 2, 1900, 426.) From water. Bacterium tiaphtJialinicus Tausson. (Planta, 4, 1927, 214.) From oil-soaked soils at Baku, Russia. Oxidizes naph- thalene . Bacterium nicolaieri Migula. (Kap- selbacillus, Nicolaier, Cent. f. Bakt., 16, 1894, 601; Migula, Syst. d. Bakt., 2, 1900, 354.) Associated with purulent nephritis. Bacterium nicutiunum Bucherer. (Cent. f. Bakt., II Abt., 105, 1942-43, 446.) From fermenting tobacco leaves. Bacterium nicotinobacter Bucherer. (Cent. f. Bakt., II Abt., 105, 1942, 170.) From a mixture of soil, manure and rot- ting materials. Gram-variable. Bacterium nicotinophagum Bucherer. (Cent. f. Bakt., II Abt., 105, 1942, 167.) From a mixture of soil, manure, and rot- ting materials. Also from fermenting tobacco leaves (ibid., 446). Bacterium nitens Kern. (Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1896, 459.) From the intestines of birds. Bacterium notuae (Sehimmelbusch) Mi- gula. (Bacillus nomae Sehimmelbusch, Deutsch. med. Wochnschr., 1889, No. 26; Migula, Syst. d. Bakt., 2, 1900, 384.) Found in necrotic tissues. Bacterium oblongum (Boutroux) De- Toni and Trevisan. (Micrococcus ob- longus Boutroux, Annales de I'Ecole normale superieure, S^r. 2, 5, 1881, 67; Bacillus oblongus Trevisan, I generi e le specie delle Batteriacee, 1889, 16; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 1021; Bacterium (jluconicum Miquel and Cambier, Traite de Bact., 1902, 605; not Bacterium glu- conicum Hermann, Biochem. Zeit., 192, 1928, 198.) From vinegar. May be an acetobacter. Bacterium ogatae Migula. (Ogata, Cent. f. Bakt., 5, 1891, 442; Migula, Syst. d. Bakt., 2, 1900, 389.) From dust. Bacterium orchiticum (Kruse) Chester. (Bacillus zur Rotzdiagnose, Kutscher, FAMILY BACTERIACEAE 683 Ztschr. f. Hyg., SI, 1895, 156; Bacillus orchiticus Kruse, in Fliigge, Die Mikro- organismen,3Aufl.,^, 1896,455; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 99.) From nasal secretions of a glandered horse. Bacterium osteophiluni Billet. (Con- tribution a r^tude de la morphologie et du developpement des Bacteriacees, Bull. Sci. de la France et de la Belgique, Paris, 31, 1890, 149.) From macerated human bones. Bacterium ovale Migula. (Bacillus No. 17, Pansini, Arch. f. pathol. Anat. u. Physiol., 122, 1890, 451; Migula, Syst. d. Bakt., 2, 1900, 458; not Bacterium ovale Chester, Man. Determ. Bact., 1901, 171 {Bacillus oralis Wright, Mem. Nat. Acad. Sci. ,7, 1895,435).) Fromfeces. Bacterium ovatum Migula. {Bacillus ovatus minutissimus Unna-Tommasoli, Monatsh. f. prakt. Dermatol., 9, 1889, 59; Migula, Syst. d. Bakt., 2, 1900, 417; Bacterium ovatus minutissimus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 139; not Bacterium ovatum Chester, Man. Determ. Bact., 1901, 177 {Bacillus ruber ovatus Bruyning, Arch, neerl. Sci. exact, et nat., Ser. II, 1898, 297).) From human skin with seborrheic eczema. Bacterium pallois Henrici. (Arb. bakt. Inst . Karlsruhe, / , Heft 1 , 1894, 36. ) From cheese. Bacterium pallescens Henrici. (Arb. bakt. Inst. Karlsruhe, 7, Heft 1, 1894, 35.) From cheese. Bacterium pallidum Henrici. (Arb. bakt. Inst. Karlsruhe, ;, Heft 1, 1894, 34; JJlvina pallida Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933, 76.) From cheese. Bacterium pallidior Chester. {Bacil- lus fuscus pallidior Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 361; Bacterium juscus pallidior Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 111; Chester, Man. Determ. Bact., 1901, 171.) Cul- ture received by Dyar from Krai's laboratory labeled Bacillus latericeus. Dyar renames this because the culture does not agree with Bacillus latericeus Eisenberg. However, the 1900 Krai catalogue indicates that this was Bacil- lus latericeus Adametz and Wichmann syn. ziegelrother Bacillus, Adametz; Bacterium lactericeum Lehmann and Neumann. Bacterium papillare Issatchenko. (Recherches sur les microbes de I'Oc^an Glacial Arctique (in Russian). Petro- grad, 1914, 149.) From sea water. Bacterium paradoxus (Kruse) Chester. (Typhus ahnlicher Bacillus, Kruse and Pasquale, Ztschr. f. Hyg., 16, 1894, 19; Bacillus paradoxus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 373; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 71.) From the liver in a case of dysentery. Bacterium paraviscosum Buchanan and Hammer. (Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 266.) Stated to be similar to Bacterium viscosum of various authors. Bacterium paleUiforme Honing. (Cent. f. Bakt., II Abt., 37, 1913, 378.) From tobacco plants in Sumatra. Bacterium pateriforme Migula. {Ba- cillus albicans pateriformis Unna-Tom- masoli, Monatsh. prakt. Dermatol., 9, 1889, 58; Migula, Syst. d. Bakt., 2, 1900, 415.) Found on human skin with seborrheic eczema. Bacterium petersii Migula. (Bac- terium C, Peters, Botan. Zeitung, 4?', 1889; Bacillus aceticus petersii Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl. 2, 1896, 355; Bacterium aceticus petersii Chester, Ann. Rept. Del. Col. Agr. Exp Sta., 9, 1897, 77; Migula, Syst. d. Bakt. 2, 1900, 397; Bacterium aceticum Chester Man. Determ. Bact., 1901, 127; not Bac- terium aceticum. Baginsky, Ztschr. f. phys. Chem., 12, 1888, 437.) From fer- menting dough. Bacterium photometricum Engelmann. (Engelmann, Jour. Roy. Microscop. Soc, 1882, 656 and 1883, 256; Bacillus photo- metricus Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) Sapro- phytic. Bacterium piluliformans (MUlIer-Thur- 684 MANUAL OF DETERMINATIVE BACTERIOLOGY gau) Migula. {Bacillus pihdiformans Miiller-Thurgau, Jahr?sber. d. Versuchs- station zu Wadenswil 1892/3, 3, 1894, 92; Migula, Syst. d. Bakt., 2, 1900, 513.) From a disease of red wine. Bacterium pituitosnm Migula. (Ba- cillus lactis pituitosi Loeffler, Berliner klin. Wochnschr., 1887, 631; Bacterium lactis pituitosi Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 86; Migula, Syst. d. Bakt., 2, 1900, 403; Bacterium lactis Chester, Man. Determ. Bact., 1901, 148; not Bacterium lactis Baginsky, Ztschr. f. phys. Chem., 12, 1888, 437.) From milk. Bacterium pityocampac DufrencA'. (Compt. rend. Soc. Biol., Paris, 71, 1919, 288.) From diseased caterpillars of the processionary moth {Cnethocnmpa piti/o- campa). Bacterium pleuropneumoniae Miguhi. (Diplococcus der Brustseuche der Pferde, Schiitz, Arch. f. pathol. Anat. u. Physiol., 707, 374; Migula, Syst. d. Bakt., 2, 1900, 348.) Frequently isolated from horses with pneumonia. Bacterium plicatum (Zinunermann) Cliester. (Bacillus plicatus Zimmer- mann, Die Bakterien unserer Trink- und Nutzwasser, Chemnitz, 1, 1890, 54; Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., ■9, 1897, 108; Bacterium plicatirum Mi- gula, Syst. d. Bakt., 2, 1900, v and 453). From water. Bacterium pneumopecurium Chester. (Bacillus of sporadic pneumonia of cattle, Smitii, V. S. Dept. Agr. Bur. Animal Husbandry, 1895, 136; Chester, Man. Determ. Bact., 1901, 137. ) Similar to Pasteurclla suillu. Bacterium pneumosepticum (Babes) Migula. {Bacillus pneumosepticus Babes, Progres m^d. roumain, 6, 1889; not Bacillus pneumosepticus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 408; Migula, Syst. d. Bakt., 2, 1900, 377.) From a case of septic pneu- monia. Bacterium polymorph um (Frankland and Frankland) Migida. (Bacillus poly- morphus Frankland and Frankland, Philos. Trans. Royal Soc. London, 178, 1887, B, 275; Migula, Syst. d. Bakt.,:?, 1900, 420.) From air. Bacterium porri Majocchi. (Majoc- chi, in Tommasi-Crudeli, Anatomia patologica, 1, 1882; Bacillus verrucae inil- garis Kuhnemann, Monatsh. f. prakt. Dermatol., 9, 1889; Bacillus porri Trevi- san, I generi e le specie delle Batteriacee, 1889, 13.) From warts. Bacterium prodeniae Metalnikov and Metalnikov. (Compt. rend. Acad. Agric, France, 18, 1932, 206.) From a blackened dead larva of a moth (Pro- denia litura) . Bacterium profusum (Frankland and Frankland) Migula. (Bacillus profusus Frankland and Frankland, Philos. Trans. Royal Soc. London, 178, 1887, B, 276; Migula, Syst. d. Bakt., 2, 1900, 421.) From air. Bacterium pseudoaquatilc Migula. (Bacillus aquatilis a, Tataroff, Inaug. Diss., Dorpat, 1891, 44; Migula, Syst. d. Bakt., 2, 1900, 470.) From water. Bacterium pseudocojij unctivitidis (Kruse) Chester. (Kartulis, Cent. f. Bakt., /, 1887, 289; Bacillus pseudncon- junctiritidis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 441; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 108.) From conjunctival secretions. Bacterium pseudofilicinum Migula. (Fadenbacillus, Maschek, Bakteriolo- gische Untersuchungen der Leitmeritzer Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 454.) From water. Bacterium pseudoinfluenzae (Kruse) Chester. (Pseudoinfluenzabacillus, Pfeiffer, Ztschr. f. Hyg., 13, 1893, 357; Bacillus pseudoinfluenzae Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 439; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 66.) From water. Bacterium pseudokeralomalaciae Mi- gula. (Loeb, Cent. f. Bakt., 10, 1891. 369; Migula, Syst. d. Bakt., 2, 1900, 359.) FAMILY BACTERIACEAE 685 A capsulated bacterium from infected cornea of a child. Bacterium psetidomultipediculum Mi- gula. {Bacillus vndlipedicidus flavus Zimmermann, Bakt. imser. Trink- u. Nutzwasser, Chemnitz, 2, 1894, 42; Mi- gula, Syst. d. Bakt., 2, 1900, 332.) From sewage. Bacterium pseudopneiimonicum (Pas- set) Chester. (Bacillus pscudopneiimo- nicus Passet, Untersuchungen iiber die Aetiologie der eiterigen Phlegmone des Menschen, 1885, 40; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 140; Brucella pscudop7^eumonicio)i Pribram, Klassifikation der Schizomyceten, Leip- zig und Wien, 1933, 68.) From pus. Bacterium punctum (Mueller) Ehren- berg. (Manas punctum ^Mueller, In- fusoria, 1786, 3; Bacillus puuctum Trevi- san, I generi e le specie delle Batteriacee, 1889, 18.) From swamps and stagnant water. Bacterium putidum Chester. (Bacillus gracilis cadaveris Sternberg, Man. of Bact., 1893, 733; Chester, :Man. Determ. Bact., 1901, 140.) From a liver. Bacterium pyacmicum INIigula . (Levy, Cent. f. klin. Med., 1890, Xo. 4; abst. in Cent. f. Bakt., 8, 1890, 86; :\Iigula, Syst. d. Bakt., 2, 1900, 443.) From a case of pyemia. Bacterium pyocinnahareum (Kruse) Chester. (Ferchmin, Ueber rote Eite- rung, Wratsch, 1892, Xo. 24 and 25; abst. in Cent. f. Bakt., 13, 1893, 103; Bacillus pyocinnahareus Kruse, in Fliigge, Die IMikroorganismen, 3 Aufl., 2, 1896, 304; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 5, 1897, 113.) From a case of red pus. Bacteriutn pyogenes Chester. (Fuchs, Inaug. Diss., Greifswald, 1890; Bacillus pyogenes anaerobius Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 244; not Bacillus pyogenes anaerohius Bela-Johan, Cent, f . Bakt., I Abt., Orig., 87, 1922, 290; Bacterium pyogenes an- aerohius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 127; Chester, Man. Determ. Bact., 1901, 184; not Bacterium pyogenes ^ligula, Syst. d. Bakt., 2, 1900, 381; not Bacterium pyo- genics Ward, Jour. Bact., 2, 1917, 619.) From stinking pus from a rabbit. Bacterium pyraustae Nos. 1-7 Metalni- kov and Chorine. (Internat. Corn Borer Invest., Sci. Repts., 1, 1928, 52.) From diseased corn borer larvae (Py- rausta nubilalis Hb.). Bacterium radiatum Chester. (Del. College Agr. Expt. Sta. Ann. Rept., 11, 1900, 56.) From soil. Bacterium ramificans Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 229.) From bean infusions. Bacterium rangiferinum Honing. (Honing, Cent. f. Bakt., II Abt., 37, 1913, 379 ; Plocamobacteri um rangiferinum Pribram, Klassifikation der Schizo- myceten, Leipzig und Wien, 1933, 78.) From fermenting tobacco. Bacterium repens ]\Iiehe. An orga- nism associated with Bacterium foliicola de Jongh. Bacterium retiformans Gicklhorn. (Cent. f. Bakt., II Abt., 50, 1920, 421.) -V sulfur bacterium from garden soil. See INIanual, 5th ed., 1939, 86 for a de- scription of this organism. Bacterium rhizopodicum Migula. (Ba- cillus rhizopodicus margarineus Jolles and Winkler, Ztschr. f. Hyg., 20, 1895, 105; Migula, Syst. d. Bakt., 2, 1900, 452.) From margarine. Bacterium roseum Losski. (Losski, Inaug. Diss., Dorpat, 1S93; quoted from ISIigula, Syst. d. Bakt., 2, 1900, 484; Bacillus roseus Xepveux, These, Fac. Pharm., Paris, 1920, 115.) From sand. Bacterium ruhigenosum Kern. (Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft, 4, 1896, 456; Bacillus ruhigenosus X'epveux, These, Fac. Pharm., Paris, 1920, 113; not Bacillus rubiginosus Catiano, in Cohn, Beitr. z. Biol. d. Pflanzen, 7, 1896, 538.) From the intestines of birds. Bacterium rubrum Schneider. (Bak- terium rubrum Schneider, Arb. bakt. Inst. Karlsruhe, 1, Heft 2, 1894, 213; also see Migula, Syst. d. Bakt., 2, 1900, 488; Bacillus ruhrum Nepveux, These, 68G M.ANUAL OF DETERMINATIVE BACTERIOLOGY Fac. Pharm., Paris, 1920, 115.) From swamp water. Difficult to distinguish from Bacterium erythromyxa. Bacterium ruhrum Metalnikov and Metalnikov. (Compt. rend. Acad. Agric, France, 18, 1932, 204; not Bac- terium, ruhrum Schneider, Arb. bact. Inst. Karlsruhe, 1, Heft 2, 1894, 213.) From the cotton worm {Gelechta gossy- piella) . Bacterium salivae Migula. (Bacillus salivae minutissimus Kruse, in Fliiggc, Die Mikroorganismen, 3 Aufl., 2, 1896, 440; Bacterium salivae minutissimus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 86; Migula, Syst. d. Bakt., 2, 1900, 418.) From secretions of the mouth. Bacterium sahuonicida Lehmann and Neumann. (Bacillus der Forellen- seuche, Emmerich and Weibel, Arch. f. Hyg., 21, 1894, 1; Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 240; Bacillus salmonicida Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 322; Bacterium sahnonica Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 99; see Mackie et al.. Final Rept. of the Furunculosis Committee, H. M. Station- ery Office, Edinburgh, 1935; and Duff, Jour. Bact., 34, 1937, 49.) Pathogenic for trout. Bacterium satiguinis Migula. (Bacil- lus sanguinis typhi Sternberg, Man. of Bact., 1893, 732; Bacterium sanguinis typhi Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 89; Migula, Syst. d. Bakt., 2, 1900, 506.) From the blood of typhus fever patients. Bacterium schujfneri Honing. (Cent, f. Bakt., II Abt., 37, 1913, 370.) From tobacco plants in Sumatra. Bacterium septentrionale Issatchenko . (Recherches sur les microbes de 1 Oc^an Glacial Arctique (in Russian). Petro- grad, 1914, 239.) From sea water. Bacterium (Proteus) septicus (Babes) Chester. (Proteus septicus Babes, Sep- tische Processe des Kindesalters, 1889; Bacillus proteus septicus Kruse, in FlUgge, Die Mikroorganismen, 3 Aufl., 2, 1896, 279; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 102; Bacillus septicus Chester, Man. Determ. Bact., 1901, 245; not Bacillus septicus Mace, Traite pratique de Bact., 1st ed., 1889, 455; not Bacillus septicus Migula, Syst. d. Bakt., 2, 1900, 646; not Bacillus septi- cus Crookshank, Textb. of Bact., 4th ed., 1900, 632.) From the intestine of a child having septicemia. Bacterium setosum Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 46.) From cheese. Bacterium siccum Issatchenko. (Re- cherches sur les microbes de I'Ocean Glacial Arctique (in Russian). Petro- grad, 1914, 235.) From sea water. Bacterium sieberti Migula. (Siebert, Inaug. Diss., Wiirzburg, 1894, 13; Migula, Syst. d. Bakt., 2, 1900, 456.) From hair follicles. Bacterium soriferum Migula. (Sev- erin. Cent. f. Bakt., II Abt., 1, 1895, 799; Migula, Syst. d. Bakt., 2, 1900, 438.) From manure. Bacterium spiniferum (Unna-Tomma- soli) Chester. (Bacillus spiniferus Unna-Tommasoli, Monatsh. f. prakt. Dermatol., 9, 1889, 58; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 110 and 143.) From human skin with seborrheic eczema. Bacterium spitwsutu Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 219.) From fermenting beets. Bacterium spirale Issatchenko. (Re- cherches sur les microbes de I'Oc^an Glacial Arctique (in Russian). Petro- grad, 1914, 238.) From sea water. Bacterium sputigenum Chester. (Ba- cillus aerogenes sputigenus capsulatus Herla, Archiv de Biol., U, 1895, 403; Chester, Man. Determ. Bact., 1901, 133; not Bacterium sputigemim Migula, Syst. d. Bakt., 2, 1900, 378.) From the blood of a mouse which had been inoculated with the sputum of a pneumonia patient. Bacterium sputigenum Migula. (Krei- bohm, Inaug. Diss., Helmstedt, 1898, 29; Migula, Syst. d. Bakt., 2, 1900, 378.) From the mouth. FAMILY BACTERIACEAE 687 Bacterium squamatum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 242.) From vegetable infusions. Bacterium squamosum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 436.) From the stomachs and intestines of birds. Bacterium stalactitigenes Honing. (Cent. f. Bakt., II Abt., S7, 1913, 375.) From tobacco plants in Sumatra. Bacterium sternbergii Migula. (Ba- cillus anaerobius liquefaciens Sternberg, Man. of Bact., 1893, 693; Migula, Syst. d. Bakt., 2, 1900, 444; Bacterium an- acrobicum Chester, Man. Determ. Bact., 1901, 198; Bacillus sternbergii Winslovv, Kligler and Rothberg, Jour. Bact., 4, 1919, 487.) From intestines of yellow fever cadavers. Bacterium steroidiclasium Arnaudi and Ercoli. (Boll. Sez. ital. Soc. intern. Microbiol., 20 (3), 1941, 000; also see Arnaudi, Cent. f. Bakt., II Abt., 105, 1942-43, 352.) Source not given in second paper. From bakers' yeast. Bacterium streckeri (Trevisan) Migula . (Bacillus citreus cadaveris Strassmann and Strecker, Ztschr. f. Medizinal- beamte, 1888, No. 3; Bacillus streckeri Trevisan, I generi e le specie delle Batteriacee, 1889, 17; Bacterium citreus cadaveris Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 108; Migula, Syst. d. Bakt., 2, 1900, 460; Bacterium citreum Chester, Man. Determ. Bact., 1901, 167.) From a cadaver. Bacterium subcitricum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 257.) From vegetable infusions. Bacterium subfuscum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 461.) From the intestines of birds. Bacterium subluteum Migula. (Bacil- lus luteus von Dobrzyniecki, Cent. f. Bakt., I Abt., 21, 1897, 835; Migula, Syst. d. Bakt., 2, 1900, 456.) From the mouth. Bacterium sulfureum Holschewnikoff. (Holschewnikoff, Fortschr. d. Med., 7, 1889, 204 and Ann. de Microgr., /, 1888- 1889, 261; Bacillus sulfureus Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From sewage. Bacterium sumatranum Honing. (Cent. f. Bakt., II Abt., 37, 1913, 374.) From tobacco plants in Sumatra. Bacterium surgeri (Dornic and Daire) Buchanan and Hammer. (Bacillus sur- geri Dornic and Daire, Bull. mens, de 1 'Office de renseignements agricoles, 6, 1907, 146; Buchanan and Hammer, Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 254.) From serum pro- duced in the manufacture of casein. Causes slimy milk. Closely related to the Bacterium bulgaricum group, accord- ing to Buchanan and Hammer. Bacterium sycosiferum Migula. (Ba- cillus sycosiferus foetides Unna-Tom- masoli, Monatsh. f. prakt. Dermatol., 8, 1889, 183; Migula, Syst. d. Bakt., 2, 1900, 385.) From the beard of a patient with bacillogenic sycosis. Bacterium syphilidis (Kruse) Migula. (Syphilisbacillus, Lustgarten, Wiener med. Wochnschr., 1884 and Wiener med. Jahrbiicher, 1885; Pacinia syphilitica Trevisan, I generi e le specie delle Batteriacee, 1889, 23; Bacillus syphilidis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 514; Migula, Syst. d. Bakt., 2, 1900, 496.) From syphilis. Bacterium tachytonum Fischer. (Fischer, Deutsche med. Wochnschr., 1894, No. 25-28; Bacillus tachy tonus Migula, Syst. d. Bakt., 2, 1900, 655.) From feces in a case of cholera. Bacterium, tenue Migula. (Bacillus tenuis sputigenes Pansini, Arch. f. path. Anat., 122, 1890, 453; Bacillus sputigenus tenuis (sic) Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 431; Bac- terium sputigenes tenuis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 89; Migula, Syst. d. Bakt., 2, 1900, 457.) Associated with advanced phthisis and catarrhal pneumonia. Bacterium termo (Mueller) Ehrenberg. (Monas termo Mueller, Infusoria, 1786; Ehrenberg, Abhandl. Akad. Berl., 1830; Palmella infusionum Ehrenberg, In- fusionsthierchen, 1838, 526; Zoogloea 688 MANUAL OF DETERMINATIVE BACTERIOLOGY iermo Cohn, Nova Acta Leop. Carol., 5^, 1853, 123; Bacillus termo Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) From infusions. Bacterium termo v;ir. subterraneinii Hansgirg. (Hansgirg, Oest. Bol . Ztschr., 1888, 6; quoted from DeToni and Trevisan, in Saccardo, Syllogo Fungorum, 8, 1889, 1025.) From damj) walls in a cellar. Bacterium thiogencs Lclmiann . (Thionsaurebakterien, Trautwein, Cent, f. Bakt., II Abt., 5S, 1921, 513; ibid., 61, 1924, 1; Lehmann, in Lehmann and Neu- mann, Bakt. Diag., 7th Aufl., 2, 1927, 516; Thiohacillus trautweinii Bergey et al . , Manual , 2nd ed . , 1925, 39. ) From canal water, sewage and soil. Regarded by Trautwein {loc. cit., 1924, 5) as closely related to Bacterium deuitrificans Leli- mann and Neumann. See Flavobac- teriuin denitrificans Bergey et al. Hetero- trophic and therefore wrongly placed in Thiobacillus (Starkey, Jour. Bact., 28, 1934, 387; Jour. Gen. Physiol., 18, 1935, 325). Bacterium tlwloeideiuii Cessner. (Gessner, Arch. f. Hyg., 9, 1889, 129; Bacillus tholoeideus DeToni and Trevi- san, in Saccardo, Sylloge Fungorum, 8. 1889, 952.) From the human duodenum . Bacterium tortuosum Zukal. (Zukal, Verb. d. zoolog. botan. Gesellsch., Wien, 35, 1885; Bacillus tortuosus Trevisan, I generi e le specie delle Batteriacee, 1889, 18; not Bacillus tortuos2is Debono, Cent, f. Bakt., I Abt., Orig., 62, 1912, 233.) From muddy water. Bacterium tremulaus Ehrenberg. (Ehrenberg, Abhandlungen d. Berliner Akad., 1830, 38; Vibrio Iremulans Ehren- berg, Die Infusionsthierchen, 1838, 79; Trevisan, Rend. 1st. Lomb., 1879, 145; Bacillus tremulans Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) From stagnant water, infusions, etc. Bacterium trichorrhexidis INIigula . {Bacillus muUiJormis trichorrhexidis Ho- dara, Monatsh. f. prakt . Dermatol., 19, 1894, 173; Migula, Syst . d. Bakt., 2, 1900, 437.) From healthy hair showing trichorrhexis. Bacterium Intncutum Chester. (Ba- cillus No. XII, Adametz, Landwirtsch . Jahrb., 18, 1889; Chester, Man. Determ. Bact., 1901, 157; not Bacterium truncatum Migula, Syst. d. Bakt., 2, 1900, 407; not Bacterium truncatum Chester, loc. cit., 195.) From Emmenthal cheese. BactcriuDi tuberosum Kern. (Arb . bakt. Inst. Karlsruhe,^, Heft 4, 1896,455; Bacillus tuberosiis Nepveux, These, Fac. Pharm., Paris, 1920, 113.) From the intestines of birds. Bacterium turcosum. (Quoted from Franke and Rudloff, Biochem. Ztschr., 310, 1942, 207.) Source not given. Bacterium uniforme Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 250.) From fermenting malt. Bacterium ureae Leube and Graser. (Leube and Graser, Arch. f. pathol. Anat. u. Physiol., 100, 1885, 558; Bacillus ureae Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 357; not Bacillus ureae Miquel, Bull. Soc. Chim. d. Paris, 31, 1879, 391; Urobacillus leubei Miciuel and Cambier, Traite de Bacteriologie, Paris, 1902, 635; not Urobacillus leubei Beijerinck, (Jent . f. Bakt., II Abt., 7, 1901, 51; Plocamo- buctcrium ureae Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 78.) From urine. Leube makes no statement regarding spore forma- tion. While Miquel's and Leube 's or- ganisms are sometimes regarded as having been identical, Miquel did not regard his Bacillus ureae as being identi- cal with Leube 's Bacterium ureae and gave them separate names Urobacillus duclauxii and Urobacillus leubei (Miquel and Cambier, loc. cit., 631 and 635). The latter name had however been pre- viously used by Beijerinck (loc. cit.) for a different organism. Dyar credits the name Bacillus ureae to Jaksch (Ztschr. f. physiol. Chem., 5, 1881, 395) who, however, spoke only of a Harnstoffpilz and evidently had no pure cultures. Dyar's culture which came from Krai is listed in the 1900 Krai catalogue as FAMILY BACTERIA CEAE 689 Bacillus ureae Leube. Also see Gibson (Jour. Bact., 24, 1935, 493). Lohnis (Handb. f. landwirtsch. Bakt., 1910, 459) thinks that this species belongs in the Proteus group. Bacterium vaillardi Migula. (Kelsch and Vaillard, Ann. Inst. Past., 4, 1890, 276; Migula, Syst. d. Bakt., ^, 1900, 437.) Found in swellings of the lymph system in leukemia. Bacterium varicosuni Migula. (Gom- bert, Recherches cxper. microbes con- jonctives, Paris, 1889; Bacillus varicosus conjunctivae Sternberg, Man. of Bact., 1893, 474; Bacterium varicosus conjunc- tivae Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 100; Migula, Syst. d. Bakt., 2, 1900, 444.) From the normal conjunctiva of man. Bacterium variosum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 218.) From vegetable infusions. Bacterium velatum Migula. (Bacillus tuberigenus 5, Gonnermann, Land- wirtsch. Jahrb., 23, 1894, 657; Migula, Syst. d. Bakt., 2, 1900, 454; Bacillus velatus Nepveux, These, Fac. Pharm., Paris, 1920, 113.) From lupine root nodules. Bacterium vernicosum Zopf. (Zopf, Beitr. z. Physiol, u. Morphol. niederer Organismen, Heft 1, 1892, 63; Bacillus rer/iicosi/s Migula, Syst. d. Bakt., 2, 1900, 781.) From cotton-seed meal. Bacterium vesiculosum Henrici. (Arb. a. d. bakt. Inst. d. techn. Hochschule zu Karlsruhe, 1, Heft, 1, 1894, 37.) From cheese. Bacterium villosum (Keck) Migula. {Bacillus villosus Keck, Inaug. Diss., Dorpat, 1890, 47; Migula, Syst. d. Bakt., 2, 1900, 429; Plucamohaclerium villosuni Pribram, Klassifikation der Schizo- myceten, Leipzig und Wien, 1933, 79.) From water. Bacterium vinicola Migula. (Bacillus viscosus vini Kramer, Bakteriol. in ihren Beziehungen z. Landwirtsch., 2, 1892, 144; Migula, Syst. d. Bakt., 2, 1900, 510.) From wine. Bacterium viyiiperda Migula. {Bacil- lus saprogenes vini IV, Kramer, Bak- teriol. in ihren Beziehungen z. Land- wirtsch., 2, 1892, 135; Migula, Syst. d. Bakt . , 2, 1900, 446.) From diseased wine . Bacterium viride van Tieghem. (Bull. Soc. bot. France, 27, 1880, 174.) Found on a fungus. Bacterium viscidum Migula. {Bacillus viscosus margarineus Jolles and Winkler, Ztschr. f. Hyg., 20, 1895, 104; Migula, Syst. d. Bakt., 2, 1900, 450.) From margarine. Bacterium viscusutu Migula. {Bacillus viscosus sacchari Kramer, Sitzungsb. d. kais. Akad. d. Wiss., Wien, 1889; Bakteriol. in ihren Beziehungen z. Landwirtsch., 2, 1892, 156; Migula, Syst. d. Bakt., 2, 1900, 447.) Similar to Leu- conostoc inesenteroides except that it liquefies gelatin. Bacterium viscusum non-liquefaciens Stutzer and Wsorow. (Cent. f. Bakt., II Abt., 71, 1927, 117.) From pupae of moth {Euxoa segetum). Resembles Ba- cillus viscosus Frankland. Bacterium vitulinum Chester. (Bacil- lus der Septikamie bei einem Seekalbe, Bosso, Cent. f. Bakt., 25, 1899, 52; Chester, Man. Determ. Bact., 1901, 143.) From a septicemia of the sea-calf {Phoca vitidina) . Bacterium vitulorum Migula. (Bacil- lus der weissen Ruhr der Kalber, Jensen, Monatsh. f. prakt. Tierheilk., 3, 1892, 92; Maanedskrift for Dyrlaeger, 4, 1892- 93, 140; Bacillus dysenteriae vitulorum Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 412; Bacterium dysenteriae vitulorum Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 86; Migula, Syst. d. Bakt., 2, 1900, 394; Bacterium dysenteriae Chester, Man. Determ. Bact., 1901, 145.) Associated with dysentery of calves. Bacterium winkleri Migula. (Mar- garinbacillus 0, Jolles and Winkler, Ztschr. f. Hyg., 20, 1895, 102; Migula, Syst. d. Bakt., 2, 1900. 485.) From margarine. Bacterium wrightii Chester. (Cap- sule Bacillus of Mallory and Wright, 690 MANUAL OF DETERMINATIVE BACTERIOLOGY Ztschr. f. Hyg., £0, 1895, 220; Chester, Man. Determ. Bact., 1901, 133.) From a case of bronchopneumonia. Bacterium zinnioides Honing. (Cent, f. Bakt., II Abt., 37, 1913, 371.) From tobacco, peanut and other plants in Sumatra. Bacterium zuernianuvi (List) Chester. {Bacillus zuernianus List, Inaug. Diss., Leipzig, 1885, 36; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 83.) From fresh manure and intestines of sheep; also found in water. Coccobacillus acridiormn Picard and Blanc. (Coccobacille des saute relics, d'Herelle, Compt. rend. Acad. Sci. Paris, 152, 1911, 1413; Picard and Blanc, ibid., 156, 1913, 1335; Bacillus acridiorum Chatton, ihid., 156, 1913, 1708.) From a locust (Schistocerca americana Drury) . Coccobacillus cajae Picard and Blanc. (Compt. rend. Acad. Sci., Paris, 156, 1913, 1334; Bacillus cajus Marchol, Re- vue de Phytopath. Appl., 1, 1914, 11.) From diseased caterpillars of Arctia caja. Coccobacillus gibsoiii Chorine. (In- ternat. Corn Borer Invest., Sci. Repts., 2, 1929, 42; B. gibsoni Paillot, B. presum- ably indicates Bacterium, see index, p. 522, L'infection chez les insectes, 1933, 134; Bacillus gibsoiii Steinhaus, Bacteria Associated Extracellularly with Insects, Minneapolis, 1942, 58.) From diseased corn borer larvae (Pyraustra nubilalis). Coccobacillus insectorum HoUande and Vernier. (Compt. rend. Acad. Sci., Paris, 171, 1920, 207.) From diseased caterpillars of a moth {Malacosoma castrensis) . Coccobacillus insectorum var. malaco- somae Hollande and Vernier. (Compt. rend. Acad. Sci., Paris, 171, 1920, 208.) From diseased caterpillars of a moth (Malacosoma castrensis) . Denitrobacterium Ihermophilum Am- broz. (Cent. f. Bakt., II Abt., 37, 1913, 3.) A thermophilic bacterium from soil. Diplobacilhis rnelolonthae Paillot. (Compt. rend. Soc. Biol., Paris, 69, 1917, 5; Annales des Epiphyties, 8, 1922, 117.) From larvae of cockchafers {Melolontha melolontha) . Diplobacillus pieris Paillot. (Annales des Epiphyties, 8, 1922, 129.) From dis- eased caterpillars of the cabbage butter- fly {Pieris brassicae) . Helicobacterium aerogenes Miller. (Deutsche Med. Wchnschr., 1^, 1886, 119; Bacillus helicoides DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 952.) From the stomach. This is the type species of the genus Helico- bacterium . Helicobacterium. klebsii Miller. (Die Mikroorganismen der Mundhohle, 2 Aufl., Leipzig, 1892, 370; quoted from Buchanan, Gen. Syst. Bact., Baltimore, 1925, 327.) From the mouth. Microbacillus citreus baregensis Robine and Hauduroy. (Compt. rend. Soc. Biol., Paris, 98, 1928, 26.) From hot sulfur springs at Bareges. Fourment (Compt. rend. Soc. Biol., Paris, 98, 1928, 588) states that this species is Bacillus luteus Fltigge, but Robine and Hauduroy (Compt. rend. Soc. Biol., Paris, 99, 1928, 317) deny this. Micrococcobacillus necroticans Pas- cheff. (See Pascheff, Bericht. d. Oph- thalmol, Gesellsch., Heidelberg, 1916, 418 or Klin. Monatsbl. f. Augenheilk., 57, 1916, 517 and 58, 1917, 97; Coccobacil- lus polymorphus necroticans, quoted from Bayer and v. Herrenschwand, Arch. f. Ophthalmol., 98, 1919, 358; Micrococco- bacillus jolymorphus necroticans Pas- cheff, Arch. d'Ophthalmol., 38, 1921, 28; Pascheff, ibid., 97.) From the human eye. Reported as the causal organism of conjunctivitis. Nitrosobacillus ihermophilus Campbell. (Sci., 75, 1932,23.) Fromsoil. Oxidizes ammonia to nitrite. Pacinia ferrarii Trevisan. (Bacillo dell' ulcera molle, Ferrari, 1885; Trev- isan, I generi e le specie delle Bat- teriacee, 1889, 23.) Pacinia fickii Trevisan. (Bacillus e des Conjunctivalsackes, Fick, 1887; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 23.) FAMILY BACTERIACEAE 691 Pacinia micheli Trevisan. (Michel, Luftstabchen des Conjunctivalsecretes, 1882; Trevisan, I generi e le specie delle Batteriacee, 1889, 23.) From the conjunctiva. Plocamohacterium acidi lactici Prib- ram. (Lange Milchsaurestfibchen, Wolff, Cent. f. Bakt., II Abt., 20, 1908, 545; Pribram, Klassifikation der Schizo- myceten, Leipzig und Wien, 1933, 76.) From milk. Plocamobacterium epidermidis (Biz- zozero) Pribram. (Leptothrix epi- dermidis Bizzozero, Arch. f. path. Anat., 98, 1896, 455; Pribram, loc. cii., 77.) From the skin. Plocamobacterium proteolyticum (Woll- man) Pribram. {Glycohacter proteolyt- icus WoUman, Ann. Inst. Past., 26, 1912, 617; Pribram, loc. oil., 118.) Plocamobacterium riibrum Pribram, loc. cit., 78. Red cheese bacterium (Kiel). Plocamobacterium tilsitense Pribram, loc. cit., 78. From Tilsit cheese (Kiel). Proteus hominis Bordoni-Uffreduzzi. (Bacterium, Bordoni-Uffreduzzi and Di Mattel, Arch, per le scienze mediche, 10, 1886, No. 7; abst. in Cent. f. Bakt., 1, 1887, 345; Bordoni-Uffreduzzi, Ztschr. f. Hyg., 3, 1888, 333; Proteus hominis capsu- latus Bordoni-Uffreduzzi, ibid.; Proteus capsulatus septicus Banti, Lo Sperimen- tale, 88; Klebsiella bordonii Trevisan, I generi e le specie delle Batteriacee, 1889, 25; Bacillus capsulatus septicus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 345; Bacterium hominis capsu- latus Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 136; Bacterium capsu- latus and Bacterium capsulatus septicus Chester, ibid., 130; Bacterium proteus Migula, Syst. d. Bakt., 2, 1900, 362; Bacterium bordonii Chester, Manual of Determ. Bact., 1901, 152.) From a case of ragpicker's disease which may have been anthrax or malignant edema. Urobacillus beijerinckii Christensen. (Christensen, Cent. f. Bakt., II Abt., 27, 1910, 357; Bacillus beijerinckii De Rossi, Microbiologia Agraria e Technica, 1927, 646.) From humus. Utilizes urea. Urobacillus jakschii Sohngen. (Sohn- gen, Cent. f. Bakt., II Abt., 23, 1909, 93; Bacillus jakschii De Rossi, Microbiologia Agraria e Technica, 1927, 646.) From garden earth. Utilizes urea. Urobacillus 7mqiieliiBeijerinck. (Cent, f. Bakt., 7, 1901, 47.) From garden earth. L5hnis (Handb. f. landwirtsch. Bakt., 1910, 459) regards this as belong- ing to the genus Proteus. Urobacillus schutzenbergii I and II ^Nliquel. (Miquel, Ann. de Micrograph., 5, 1893, 321 and 323; Bacillus schutzen- bergii Migula, Syst. d. Bakt., 2, 1900, 727.) From sewage and river water. These may belong to Proteus (Lohnis, Handb. f. landwirtsch. Bakt., 1910, 459). Urobacterium aerophilum Rubentschik. (Cent. f. Bakt., II Abt., 66, 1925, 175.) From salt water. Lake Liman near Odessa. Urobacterium citrophilum. Rubent- schik. (Cent. f. Bakt., II Abt., 6^, 1925, 172.) From black mud and salt water, Lake Liman near Odessa. Viscobacterium laclis foetidum Laxa. (Cent. f. Bakt., II Abt., 95, 1936, 130.) From milk having a fetid odor. 692 MANUAL OF DETERMINATIVE B.VCTERIOLOGY APPENDIX TO SUBORDER EUBACTERIINEAE Record of species and synonyms discovered too late to be entered in the main body of the text. Arranged alphabetically by genera. Acetobacter aceti (Kiitzing) Beijerinck syn. Bacterium aceticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., S, 1897, 77. Acetobacter acetosum Bergey et al. syn. Ulvina acetosa Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 76. Acetobacter ascendens Bergey et al. syn. Ulvina ascendens Pribram, Klassifi- kation der Schizomyceten, Leipzig und Wien, 1933, 75. Acetobacter diversion Ilumm. (Duke Univ. Marine Lab., North Carolina, Bull. 3, 1946, 03.) From sea water, Beaufort, North Carolina and marine algae, Miami, Florida. Digests agar. Acetobacter mobile T6sic and Walker. (Jour, of Brewing, 50, 1944, 296.) From bottled ale. Acetobacter pasteurianiDn (Hansen) Beijerinck syn. Bacillus pasleurianum Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 314; not Bacillus pasteurianus Leh- mann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 82; Ulvina pasteuriana Pribram, Klassifikation der Schizomy- ceten, Leipzig und Wien, 1933, 76. Acetobacter potens Humm. (Duke Univ. Marine Lab., North Carolina, Bull. 3, 1946, 63.) From intertidal sand, Beaufort, North Carolina. Digests agar. Acetobacter rancens Beijerinck syn. Ulvina rancens Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 76. Acetobacter singulare Humm. (Duke Univ. Marine Lab., North Carolina, Bull. 3, 1946, 62.) From sea water, Beaufort, North Carolina. Digests agar. Acetobacter xylinum (Brown) Holland syn. Bacillus xylinus Trevisan, I generi e le specie delle Batteriacee, 1889, 16; Ulvina xylina Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 76. Achromobacter caseinicuui Gahl. (Jour. Bact., 16, 1928, 38.) From a solution of sodium caseinate. Polar flagellate. Possibly a strain of Pseudo- monas fluorescens Migula that had lost the power of forming pigment. Achromobacter nijibetsui Takeda. (Cent. f. Bakt., II Abt., 94, 1936, 48.) From diseased salmon eggs. Not found to be virulent. A polar-flagellated, Gram-negative, yellow chromogen, pre- sumably belonging in the genus Xan- tlwmonas. Actinobacillus actinomycetemcomitans Topley and Wilson syn. Bacillus acti- nomycetemcomitans Rosebury, Bact. Rev., 8, 1944, 205. Aerobacter liquefaciens Beijerinck. (Cent. f. Bakt., II Abt., 6', 1900, 199; not Aerobacter liquefaciens Grimes and Hennerty, Sci. Proc. Roy. Dublin Soc, (N.S.) W, 1931, 93.) From mud and water in swamps. Monotrichous, other- wise like Aerobacter cloacae. This may have been a species of gas-forming Pseudomonaa. Aerobacter tarlarivorum Nijdam. (Thesis, Leiden, 1907.) Decomposes d- tartrates. Probably identical with Aerobacter aerogencs (Vaughn, Marsh, Stadtman and Cantino, Jour. Bact., 52, 1946, 324). Alcaligenes marshallii Bergey et al. syn. Bacterium viarshaUi Buchanan and Hammer, Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 272. Alcaligenes viscosuni Weldin syn. Plo- camobacterium viscosum Pribram, Klassi- fikation der Schizomyceten, Leipzig und Wien, 1933, 79. Ascncoccus buccalis Miller. (Die Mi- kroorganismen der Mundhohle, Leipzig, 1889, 65.) From the mouth. APPENDIX TO SUBORDER EUBACTERIINEAE 693 Bacillus annulatus Wright Syn. Bac- terium annulatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 105. Bacillus cuhonianus Macchiati syn. Bacterium cubonianus Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 132. Bacillus duplicatiis Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 457; Bacterium duplicatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 90.) From Schuylkill River water. Monotrichous. Bacilhis fluorescens mutabilis Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 449; Bacterium fluorescens mutabilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 120.) From Schuylkill River water. Bacillus fluorescens nivalis Eisenberg syn. Bacterium fluorescens nivalis Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 120. Bacillus hayducki Henneberg syn. Plocamohacterium hayducki Pribram, Klassifikation der Schizomyceten, Leip- zig und Wien, 1933, 77. Bacillus influenzoidcf! apis White. (Jour. Path, and Bact., U, 1921, 71.) From intestine of bee. Monotrichous. Bacillus mesentericus aureus Winkler syn. Bacillus winhleri Chester, Man. Determ. Bact., 1901, 256. Bacillus pabuli acidi II Weiss syn. Plocamohacterium pabuli Pribram, Klas- sifikation der Schizomyceten, Leipzig und Wien, 1933, 78. Bacillus vaginae Kruse syn. Bacterium vaginae Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 67. Bacillus viridi-luteus Trevisan. (Griingelber Bacillus, Eisenberg, Bakt. Diag., 1 Aufl., 1886, 10; Trevisan, I generi e le specie delle Batteriacee, 1889, 19.) From water. This probably was the same as Bacillus fluorescens Trevisan, ibid., 19 and Pseudomonas fluorescens Migula. Bacillus wortmannii Henneberg syn. Plocamobacterium wortmanni Pribram, Klassifikation der Schizomyceten, Leip- zig und Wien, 1933, 79. Bacterium granulosum Lehmann and Neumann syn. Plocamohacterium granu- losum Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 77. Bacterium lipolyticum Huss syn. Kur- thia lipolyticum Pribram, Klassifikation der Schizomyceten, Leipzig und Wien, 1933, 75. Bacterium orleanense Henneberg syn. Ulvina orleanensis Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933, 75. Bacterium xylinoides Henneberg syn. Ulvina xylinoides Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933, 76. Brucella byzantinea (Montsouris) Prib- ram. (Coccobacterium hyzantineum Montsouris, quoted from Pribram, Klas- sifikation der Schizomyceten, Leipzig und Wien, 1933, 67; Pribram, idem.) Brucella cocciformis (Jaiser) Pribram. (Bacterium cocci forme Jai.ser, quoted from Pribram, Klassifikation der Schizo- myceten, 1933, 67; Coccobacterium ther- mophilum Negre, Compt. rend. Soc. Biol., Paris. 75, 1913, 814 and 867; Prib- ram, idem.) From sputum. Chlorobacterium lactis Guillebeau. (Landw. Jahrb. d. Schweiz., 4, 1890, 32.) From the udder of cows with mastitis. Produces a green pigment . Presumably identical with Pseudomonas aeruginosa Migula. The type species of the genus Chlorobacterium Guillebeau. Chromobacterium chocolatum Knutsen. (Quoted from Lasseur, Dupaix-Lasseur and Melcion, Travaux du Lab. de Micro- biol., Fac. Pharm. de Nancy, Fasc. XIII, 1942-43-44, 1944, 164, 187, 293, and 313.) Isolated by M. H. Knutsen, State Coll., Pennsylvania. Source not known. Dull violet with brown tinge. Disso- 694 MANUAL OF DETERMINATIVE BACTERIOLOGY ciates into a violet and an orange strain (Chromobacterhnn orangium Knutsen, loc. cil., 294). Chromobacterium iodinuni Davis. (Davis, Cent, f . Bakt., II Abt., 100, 1939, 273; also see Clemo and Mcllwain, Jour. Cham. Soc.,Pt. 1,1938, 479; Psewdowonas iodinum Tobie and Pseudomonas clemo Tobie, Bull. Assoc, des Diplomes de Mi- crobiol., Fac. Pharm. Nancy, Xo. 18, 1939, 16.) From plate inoculated with milk. This non-motile organism does not have the characters of Chromobac- terium sensu stricto so that this species is retained with Bacterium for the present . Coccus cumulus minor Black. (Trans. 111. State Dental Soc, 22, 1886, 192.) From the mouth. Corynebacterium hemolyticum Mac- Lean, Liebow and Rosenberg. (Jour. Inf. Dis., 79, 1946, 69.) From infections among American soldiers and natives in the South and West Pacific. Similar in many ways to Corynebacterium pyo- genes and C. oris. ' Corynebacterium piriformc Honing. (Cent', f. Bakt., II Abt., 37, 1913, 383.) From tobacco plants in Sumatra. Diplococcus aquatilis Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 184.) From water. Diplococcus glycinophilus Cardon and Barker. (Jour. Bact., 52, 1946, 629.) From marine mud. Diplococcus luteus Adametz and Wich- mann. (Adametz and Wichmann, Die Bakterien der Trink- und Nutzwasser, Mitt. Oest. Versuchsstat. f. Brauerei u. Malzerei, Heft 1, 1888, 49; Planococcus hiteus Migula, Syst. Bakt., 2, 1900, 274.) From water. Escherichia Castellani and Chalmers syn. Colibacter Pestana and Andrade, Ann. Paulistas de Med. e Cir., 39, 1940, 462. Escherichia coli Castellani and Chal- mers syn. Colibacter commune Pestana and Andrade, loc. cit. Flavobacterium harrisonii Bergey et al. syn. Bacillus harrisonii Buchanan and Hammer, Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 257. Flavobacterium tabidum Kimata. (Cent. f. Bakt., II Abt., 105, 1942, 120.) From spoiled semi-dried fish {Trachurus japonicus). Polar flagellate. Fusiformis grandis Grasse. (Compt. rend. Soc. Biol., Paris, 9A, 1926, 1014; Arch. Zool. Exper. et Gen., 65, 1926, 463.) From the surface of the body of a flagellate (Polymastix mclolonthae) , in the intestine of larvae of beetles and tipulids, possibly also free in the in- testine of the insects. Fusiformis legcri Grasse. (Compt. rend. Soc. Biol., Paris, 94, 1026, 1014; Arch. Zool. Exper. et Gen., 65, 1926, 467.) From the surface of the body of a flagellate {Polymastix legeri) and in the intestine of diplopods. Fusiformis lophomonadis Grasse. (Compt. rend. Soc. Biol., Paris, 9^, 1926, 1015; Arch. Zool. Exp^r. et G^n., 65, 1926, 468.) From the surface of the body of a flagellate {Lophomonas striata) and in the intestine of cockroaches. Fusiformis melolonthae Grasse. (Compt. rend. Soc. Biol., Paris, 54, 1926, 1014; Arch. Zool. Exp^r. et G^n., 65, 1926, 465.) From the surface of the body of a flagellate (Polymastix melo- lonthae) and in the intestine of larvae of beetles and tipulids. Gluconoacetobacter cerinus Takahashi and Asai. (Cent. f. Bakt., II Abt., 93, 1936, 252.) From fruits. Gluconoacetobacter liquefacicns Taka- hashi and Asai, loc. cit. From fruits. Gluconoacetobacter roseus Takahashi and Asai. (Bacterium industrium var. hoshigal-i Takahashi and Asai, Cent. f. Bakt., II Abt., 82, 1930, 400; Bacterium hoshigaki var. glucuronicum I Takahashi and Asai, ibid., 87, 1933, 385; Takahashi and Asai, ibid., 93, 1936, 252.) From dried persimmons (hoshigaki). Gluconobaclcr liquefaciens Asai. (Jour. Agr. Chem. Soc. Japan, 10, 1934, APPENDIX TO SUBORDER EUBACTERIINEAE 695 621 and 11, 1935, 50; see Cent. f. Bakt., II Abt., 93, 1936, 248.) From fruits. Jodococcns magnus Miller. (Deutsche med. Wchnschr., 14, 1888, 612.) From the mouth. The type species of the genus Jodococcns (syn. lodococcus) Miller. Jodococcns parvus ^liller {ibid., 612). From the mouth. Lactobacillus bulgaricus Holland syn. Bacterium bulgaricum Buchanan and Hammer, Iowa Sta. Agr. Exp. Sta., Res. Bull. 22, 1915, 250. Lactobacillus buchneri Bergey et al. syn. Ulvina buchneri Pribram, Klassifi- kation der Schizomyceten, Leipzig und Wien, 1933, 75. Lactobacillus delbrueckii Beijerinck syn. Ulvina delbruecki Pribram, loc. cit., 75; Plocamobaclerium delbruecki Prib- ram, ibid., 77. Lactobacillus helveticus Holland syn. Plocamobaclerium casei Pribram, loc. cit., 77; Plocamobacterium helveticum Pribram, ibid., 78. Lactobacillus pastorianus Bergey et al. syn. Plocamobacterium pastorianum Prib- ram, loc. cit., 78. Lactobacillus pentvaceticus Fred, Pe- terson and Davenport sj^n. Plocamo- bacterium pcnloaccticum Pribram, loc. cit., 78. Lactobacillus plantarum Holland syn. Ulvina cucumeris fermentati Pribram, loc. cit., 75. Lactobacillus taette Olsen-Sopp. (Cent. f. Bakt., II Abt., 33. 1912, 14.) From ropy milk. Leptotrichia Trevisan partial sj'n. Leucothrix Oersted, De regionibus marinis, 1844, 44. Listeria monocytogenes Pirie syn. Bru- cella monocytogenes Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933, 68. Mammococcus gorini. (Quoted from L. Gorini, Enzymologia, 10, 1942, 102.) From the udder. Micrococcus afermentans Castellani. (Proc. Soc. Exp. Biol, and Med., 25, 1928, 536; also see Jour. Trop. Med. and Hyg., 35, 1932, 372.) From an ulcerative lesion of the skin. Micrococciis albus var. maltigenes Du- mais and Albert. (Quebec Laitier, 5 (2), 1946, 19.) From Richelieu cheese. Regarded as an important ripening agent. Micrococcus aquatiiis Vaughan. (Vaughan, Amer. Jour. Med. Sci., 104, 1892, 190; not Micrococcus aquatiiis Chester, Man. Determ. Bact., 1901, 88; not Micrococcus aquatiiis Bolton, Ztschr. f. Hyg., 1, 1886, 94.) From water. Micrococcus aquatiiis albissimus von Rigler. (Hyg. Rund., 12, 1902, 482.) From bottled mineral waters. Micrococcus aquatiiis albus Vaughan. (Vaughan, Amer. Jour. Med. Sci., 104, 1892, 182; not Micrococcus aquatiiis albus Toporoff, Cent. f. Bakt., 13, 1893, 487.) From water. Micrococcus aquatiiis magnus Vaughan . (Amer. Jour. Med. Sci., 104, 1892, 182.) From water. Micrococcus aquivivus ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 275.) From sea water. Micrococcus cyaneus (Schroeter) Cohn syn. Bacterium cyaneus White, U. S. D.A., Bur. Entomol. Tech. Ser. Bull. 14, 1906, 16. Micrococcus enteroideiis Castellani. (Proc. Soc. Exp. Biol, and Med., 25, 1928, 536; also see Jour. Trop. Med. and Hyg., 35, 1932, 372.) From feces. Micrococcus euryhalis ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 255.) From sea water. Micrococcus griseus Winter sj'n. Ba- cillus griseus Trevisan, I generi e le specie delle Batteriacee, 1889, 18. Micrococcus himonoi Kimata. (Cent, f. Bakt.. II Abt., 105, 1942, 116.) From spoiled semi-dried fishes (Scotnber ja- ponicus and Trachurus japonicus) . Re- sembles Micrococcris caseolyticus and M. mucofaciens. 696 MANUAL OF DETERMINATIVP: BACTERIOLOGY Micrococcus infimus ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 262.) From marine bottom deposits. Micrococcus laevulosinertis Castellani. (Proc. Soc. Exp. Biol, and Med., ^5, 1928, 536; also see Jour. Trop. Med. and Hyg., 35, 1932, 372.) From a case of stoma- titis. Micrococcus maripuniceus ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 264.) Sessile form found on slides submerged in sea water. Micrococcus metentericus Castellani. (Quoted from Jour. Trop. Med. and Hyg., 35, 1932, 372.) From case of ulcerative colitis. Micrococcus moricolor Holmes and Wilson. (Jour. Bact., 49, 1945, 311.) From contaminated wounds. Produces a mulberry pigment on potato. Micrococcus myceticus Castellani. (Arch. Dermat. and Syphil., 18, 1928, 857.) From cases of pseudomycosis. Micrococcus nexifcr Miller. (Miller, Die Mikroorganismen der Mundhohle, Leipzig, 1889, 65.) From the mouth. Probably Streptococcus brevis according to Goadby (Mycology of the Mouth, London, 1903, 60). Micrococcus pvtatus Ravenel. (Mem. Nat. Acad. Sci., 8, 1896, 21.) From soil. Micrococcus putneus Castellani. (Quoted from Jour. Trop. Med. and Hyg., 35, 1932, 372.) From a case of glossitis. Micrococcus rhodochroiis Migula syn. Bacillus rhodochrous Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 362; Bacterium rhodo- chrous Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 116.) Dyar had the original Micrococcus rhodochrous culture from Krai and felt as have others who have examined this culture that it is not a true Micrococcus. Micrococcus sedentarius ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 260.) Sessile form found on slides submerged in sea water. Micrococcus sedimenteus ZoBell and Upham. (Bull. Scripps Inst, of Ocean- (jgraphy, Univ. Calif., 5, 1944, 265.) Sessile form found on slides submerged in sea water and in marine mud. Micrococcus visicidus Castellani. (Proc. Soc. Exp. Biol, and Med., 25, 1928, 536; also see Jour. Trop. Med. and Hyg., 35, 1932, 372.) From an in- flamed upper lip. Microspira vacillans Gicklhorn. (Cent. f. Bakt., II Abt., 50, 1920, 422.) From the pool in the Botanical Garden, Univ. Graz, Austria. Contains grains of sulfur. Neisseria babesi Trevisan. (Bact^rie de I'hemoglobinurie du boeuf. Babes, 1888; Trevisan, I generi e le specie delle Batteriacee, 1889, 32.) Neisseria lulea (Adametz) Trevisan. {Diplococcus luteus Adametz, 1887; Trev- isan. I generi e le specie delle Batteriacee, 1889, 32.) Neisseria michcli Trevisan. (Tra- chomcoccus, Michel, 1886; Trevisan, I generi e le specie delle Batteriacee, 1889, 32.) Neisseria pharyngis syn. Micrococcus pharyngis Cruikshank and Cruikshank, iVIed. Res. Council Syst. of Bact., 8, 1931, 349. Pacinia decipiens Trevisan. {Spiril- lum aus der Luft, Babes, Ztschr. f. Hyg., 5, 1888, 183; Trevisan, I generi e le specie delle Batteriacee, 18S9, 24.) From the air. Pacinia rabida Trevisan. (Spirillum bei Rabies, Babes, Ztschr. f. Hyg., 5, 1888, 181; Trevisan, I generi e le specie delle Batteriacee, 1889, 23.) Pectobacterium delphinii Waldee. (Ark, Phytopath., 38, 1938, 281; Waldee, Iowa State Coll. Jour. Sci., 19, 1945, 471.) Causes larkspur bacterial blight. Phytomonas asplenii Ark and Tomp- kins. (Phytopath., 36, 1946, 760.) Causes leaf blight of bird's nest fern. Phytom onas mac ulifc li um-gardeniae APPENDIX TO SUBORDER EUBACTERIINEAE 697 Ark. (Phytopath., 36, 1946, 867.) From gardenia {Gardenia jasminoides). A xanthomonad. Phytomonas syringae populates Smith. (Jour. Agr. Res., 68, 194-4, 269.) Con- sidered the cause of blister spot, a disease of apple. Phytomonas washingtoniae Pine. (Phytopath., 33, 1943, 1203.) From the Washington pahn, Washingtonia filifera. A pseudomonad. Pneumococcus flavescens Arloing. (Compt. rend. Acad. Sci., 109, 1889, 428 and 459.) From lesions of cattle having peripneumonia. Pneumococcus gutia cerei ivrloing, loc. cit. From lesions of cattle having peripneumonia. Pneumococcus lichnoides Arloing, loc. cit. From lesions of cattle having peripneumonia. Pseiulomonas aesiuniarina ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 269.) A marine sedentarj- organism. Pscudomonas allii (Griffiths) Aligula syn. Bacillus allii Sternberg, Man. ot Bact., 1893, 629. Pseudomonas ambigua (Wright) Ches- ter syn. Bacterium ambiguus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 71. Pseudomonas atlantica Humm. (Duke Univ. Marine Lab., North Carolina, Bull. 3, 1946, 58.) From seaweed (Graci- laria hlodgettii) and beach sand. Digests agar. Pseudomonas aurea Aligula syn. Bac- terium fluorescens aureus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 109. Pseudomonas azotogena ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 260.) From sea water and marine mud. Pseudomonas beaufortensis Humm (loc. cit., 58). From seawater, bottom mud and on algae. Digests agar. Pseudomonas berolinensis Migula. (Indigoblauer Bacillus, Claessen, Cent. f. Bakt., 7, 1890, 13; Bacillus berolinensis indicus Germano, Cent. f. Bakt., 12, 1892, 517; Bacillus indigoferus Zimmer- mann, Bakt. unserer Trink- u. Xutz- wasser, Chemnitz, 2, 1894, 16, not indi- cated as being the same as Bacillus indi- goferus Voges, Cent. f. Bakt., H, 1893, 307; Bacillus indigonaceus Schneider, Arb. bakt. Inst. Karlsruhe, 1, Heft 2, 1894, 228; Migula, in Engler and Prantl, Die natiirl. Pflanzenfam., 1, la, 1895, 29; Bacterium, indigonaceum Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 267; Bacterium berolinensis indicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 118.) From Spree River water. Pseudomonas butyri Migula syn. Bac- terium butyri fluorescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 120. Pseudomonas centrifugans (Wright) Chester. (Bacillus centrifugans Wright, Mem. Nat. Acad. Sci., 7, 1895, 462; Bac- terium centrifugans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 95; Chester, Man. Determ. Bact., 1901. 312.) From water. Pseudomonas coadunata Chester syn. Bacterium coadunalus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 90. Pseudomonas coenobios ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 272.) From film of marine fouling organisms. Pseudomonas cohaerea (Wright) Ches- ter, not Bacillus cohaerens Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 458; Bacterium cohaereus (sic) Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 93. Pseudomonas coli communis Conn, Esten and Stocking. (Storrs Agri. Exp. Sta., Conn., 18th Ann. Rept. for 1906, 186.) From cheddar cheese. Like Bacillus coli communis except that it has a single, long flagellum. Pseudomonas convexa Chester syn. Bacterium fluorescens convexus Chester, 698 MANUAL OF DETERMINATIVE BACTERIOLOGY Ann. Rept. Del. Col. Aj^r. Exp. Sta., 9, 1897, 123. Pseudomonas corallina Humm (loc. cit., 59). From marine algae of all conamon species at Beaufort, Nor. Car. Digests agar. Pseudomonas delabens (Wright) Ches- ter syn. Bacterium delabens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 97. Pseudomonas eisenhergii Migula syn. Bacterium fluorescens non-Uquefaciens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas elongata Humm {loc. cit., 60). From intertidal sand, Atlantic Beach, Nor. Car. Digests agar. Pseudomonas enalia ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 254.) From sea water and marine mud. Pseudomonas fairmountensis (Wright) Chester syn. Bacterium fairmountensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 90. Pseudomonas felthami ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 267.) From marine mud. Pseudomonas jimhriata (Wright) Chester syn. Bacterium fimbriaius Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 95. Pseudomonas floridana. Hunun {loc. cit., 60). From algae and beach sand at Miami, Fla., and Beaufort, Nor. Car. Digests agar. Pseudomonas fluorescens Migula syn. Bacterium fluorescens liquefaciens Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 120. Pseudomonas foliacea Chester syn. Bacterium fluorescens foliaceus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 122. Pseudomonas geniculatus (Wright) Chester. Bacillus geniculatus Wright; not Bacillus geniculatus DeBary, Inaug. Diss., Strassburg, Leipzig, 1885; not Bacillus geniculatus Trevisan, I generi e le specie delle Batteriacee, 1889, 16; syn. Bacterium geniculatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 95. Pseudomonas humicola Bersteyn. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 97.) From soil. Pseudomonas hypothermis ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 276.) From marine bottom deposits. Pseudomonas incognita Chester syn. Bacterium fluorescens incognitus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 122. Pseudomonas indoloxidans Gray. (Proc. Roy. Soc. London, B, 102, 1928, 263.) From soil from Italian Tyrol. Pseudomonas indigoferus (Voges) Mi- gula. (Bacillus indigoferus Voges, Cent, f. Bakt., IJ^, 1893, 307; Bacterium indigo- ferus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 118; Migula, Syst. d. Bakt., 2, 1900, 950.) From Kiel tap water (Voges); from Delft ditch water, mud and garden soil (Elazari-Volcani, Arch. f. Mikrobiol., iO, 1939, 357). Some authors regard Voges' organism as identical with Claessen's indigo blue bacillus, see Pseudomonas berolinensis. Pseudomonas indigoferus var. immo- bilis Elazari-Volcani. (Arch. f. Mikro- biol., 10, 1939, 350.) From ditch mud. See Lehmann and Neumann (Bakt. Diag., 1 Aufl., 2, 1896, 267) who also had a non-motile strain {Bacterium indigo- naceum) from Krai which they con- sidered identical with Claessen's indigo blue bacillus. Pseudomonas inertia Humm (loc. cit., 61). From intertidal sand, Atlantic Beach, Nor. Car. Digests agar. Pseudomonas iris Migula syn. Bacil- lus fluorescens crassus Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 294; Bacterium fluorescens crassus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 134; Bacterium iris Chester, idem, 137.) From sputum. Pseudomonas jaegeri Migula S3'n. Bac- APPENDIX TO SUBORDER EUBACTEEIINEAE 699 terium proteus fluorescens Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 119; and Bacillus urinae Chester, Man. Determ. Bact., 1901, 263. Pscudomonas javanica (Eijkmann) Mi- gula syn. Bacterium javanicnsis Ches- ter, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 111. Pseudomonas liquida Chester. {Ba- cillus liquidus Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 382; Chester, Man. Determ. Bact., 1901, 'ill; Achromo- bacter liquidum Bergey et al.. Manual, 1st ed., 1923, 14o.) From water. Orig- inally described merely as motile; Ches- ter recognizes the species as polar flagellate and lists Bacillus Uquefaciens cotmnunis Sternberg and Bacillus aqua- tilis communis Kruse as synonyms. Pseudomonas longa Migula syn. Bac- terium fluorescens longus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas macroselmis ^Migula sj'n. Bacillus fluorescens putidus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas marinopersica ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 275.) From marine bottom deposits. Pseudomonas melochlora (Winkler and Schrotter) Migula syn. Bacterium melo- chlorus Chester, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 120; see abst. in Cent. f. Bakt., 9, 1891, 700. Pseudomonas memhranula ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 270.) Sessile form found on slide submerged in sea. Pseudomonas miiuttissima ^Migula syn. Bacterium fluorescens minutissimus Ches- ter, Ann. Rept. Del. Col. Agr. E.xp. Sta., 9, 1897, 120. Pseudomonas monadijormis (Kruse) Chester syn. Bacterium monadiformis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 69; Bacterium coli mobilis Chester, ibid., 69. Pseudomonas muUistriata (Wright) Chester syn. Bacterium multistriatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 90. Pseudomonas nebulosa (Wright) Ches- ter syn. Bacterium nebulosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 93. Pseudomonas neritica ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 255.) From sea water and marine mud. Pseudomonas nexibilis (Wright) Ches- ter sj'n. Bacterium nexibilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74. Pseudomonas obscura ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 274.) From marine bottom deposits. Pseudomonas oceanica ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 266.) From marine mud. Pseudomonas ochracea (Zimniermann) Chester sj'n. Bacterium ochraceus Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 104. Pseudomonas oralis Chester syn. Bac- terium fluorescens oralis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 123. Pseudomonas pallescens Migula syn. Bacterium riridis pallescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas perjectomarinus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 277.) From sea water and marine mud. Pseudomonas periphyta ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 276.) Sessile form found in film of marine fouling organisms. Pseudomonas phospliorescens (Fisher) Bergey et al. syn. Pasteurella phos- phorescens Trevisan, I generi e le specie delle Batteriacee, 1889, 21; Bacillus phos- phorescens indicus Eisenberg, Bakt. 700 MANUAL OF DETERMINATIVE BACTERIOLOGY Diag., 3 Aufl., 1891, 123; Vibrio indicus Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 341; Bacterium phos- phorescens indicus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 120; Photobacter indicum Beijerinck, Proc. Sect. Sci., Kon. Akad. v. Wetensch., Amsterdam, 3, 1900, 352; Microspira phosphorescens Chester, Man. Determ. Bact., 1901, 333; Bacillus indicus Bei- jerinck, Folia Mikrobiologica, Delft, I, 1912, 1. Beijerinck (loc. cit., 1900) dis- cusses two variants of this species: Photobacter indicum var. obscurum and Photobacter indicum var. parvum. Later, Beijerinck {loc. ci7., 1912), in dis- cussing mutants of this species, proposes the species names Bacillus indicus par- vus, Bacillus indicus semiobscurus and Bacillus indicus obscurus. Pseudomonas piscova Hanzawa and Takeda. (Jozognku Zasshi, Osaka, Ja- pan (Jour, of Zymology), 9, 1931, 571; quoted from Takeda, Cent. f. Bakt., II Abt., 94, 1936, 46.) From diseased sal- mon eggs. Pseudomonas pleuniurpha ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 275.) From marine bottom deposits. Pseudomonas pullulans (Wright) Chester syn. Bacterium pullulans Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 105. Pseudomonas puris Patrick and Werk- man. (Proc. Iowa Acad. Sci., 37, 1930, 57.) From a typhoid-like infection of snakes. Pseudomonas ribicola Bohn. (Jour. Agr. Res., 73, 1946, 288.) From the na- tive currant, Ribes aureum. Pseudomonas riboflavinus Foster. (Jour. Bact., 47, 1944, 30.) Oxidizes riboflavin to lumichrome. From ribo- flavin-rich soil. Pseudomonas roseola Ilumm (loc. cit., 62). From intertidal sand, Atlantic Beach, Nor. Car. Digests agar. Pseudomonas rugosa (Wright) Chester syn. Bacferiuru rugosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 122. Pseudomonas schuylkilliensis Chester syn. Bacterium fluorescens schuylkil- liensis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 119. Pseudomonas scissa (Frankland and Frankland) Migula syn. Bacterium scis- sus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 143. Pseudomonas sessilis ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 259.) Sessile form found on solid surfaces sub- merged in the sea. Pseudomonas sinuosa (Wright) Chester syn. Bacterium sinuosiis Chester, Ann. liept. Del. Col. Agr. Exp. Sta., 9, 1897, 69. Pseudomonas sniaragdina Migula syn. Bacterium smaragdino foetidus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 119; and Bacillus smaragdinus Chester, Man. Determ. Bact., 1901, 263. Pseudomonas sterotropis ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 272.) From a film of marine fouling organisms. Pseudomonas striata Chester syn. Bacterium striatus viridis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 123. Pseudomonas syncyanea Migula syn. Bacterium syncyanus (sic) Schroeter, Beitr. z. Biol. d. Pflanz., 1, Heft 2, 1872, 126 and Bacterium cyanogenum Zopf , Die Spaltpilze, 2 Aufl., 1884, 50; may be in 1 Aufl. Pseudomonas synxantha (Ehrenberg) Holland. {Vibrio synxanthus Ehren- berg, Verhandl. d. Berl. Akad., 1840, 202; Vibrio xanthogenes Fuchs, Magazin f. d. ges. Tierheilk., I, 1841, 193; Bacterium xanthinum Schroeter, in Cohn, Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 120; Bacillus synxanthus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 290; Flavobac- terium synxanthum Bergey et al.. Man- ual, 1st ed., 1923, 102; Holland, Jour. Bact., 5, 1920, 220.) Bergey et al. APPENDIX TO SUBORDER EUBACTERIINEAE 701 (Manual, 1st ed., 1923, 102) give Bac- terium xanthogenes as a synonym. From milk and cream. Polar flagellate (Ham- mer, personal communication). See Hammer, Res. Bui. 20, Iowa Agr. Exp. Sta., 1915, for a description of this orga- nism. Pseudomonas tenuis Migula syn. Bac- teriimi flitorescens tenuis Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas turcosa Migula syn. Ttirkisfarbener5aa7Z«s, Tataroff, Inaug. Diss., Dorpat, 1891, 52. Pseudomonas vadosa ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 263.) From sea water and marine bottom deposits. Pseudomonas rendrelli Tobie. {Pseu- domonas vendralli {sic), mentioned by Farrell and Wolff, Jour. Ind. and Eng. Chem., 33, 1941, 1186; U. S. Patent 2,227,716, Mar. 1942, issued to Lockwood et al.; Bacillus vendrelli Lasseur, Du- paix-Lasseur and Melcion, Travaux Lab. Microbiol. Fac. Pharm. Nancy, Fasc. 13, 1942-43-44, 1944, 293.) Isolated by W. C. Tobie in 1938 from well near Ponce, Puerto Rico owned by Mr. Ven- drell (Tobie, Jour. Bact., 52, 1946, 685). Presumably Pseudomonas aeruginosa. Pseudomonas virescens (Frick) Migula syn. Bacterium virescens Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 124. Pseudomonas viscosa (Frankland and Frankland) Migula syn. Bacterium vis- cosus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 145; not Bac- terium viscosum Weldin and Levine, Abst. Bact., 7, 1923, 16. Pseudomonas xanthochrus ZoBell and Upham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 279.) From marine bottom deposits. Ramibacterium alactolyticum Pr^vot and Taffanel. (Ann. Inst. Past., 68, 1942, 259.) From an osteoplilegmon of the maxillarv bone. Salmonella atherton Ferris, Hertzberg and Atkinson. (Med. Jour. Australia, 32, 1945, 368.) From 29 cases of gastro- enteritis in an army hospital. Salmonella typhosa (Zopf) White syn. Bacillus typhicus Cabral and Da Rocha, 1. Trabalhos do Cabinette de Micro- biologia; abst. in Ann. de Micrographie, 2, 1889-1890, 295. Sarcina pelagia ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 279.) From sea water and marine bottom deposits. Serratia fuchsina Bergey et al. syn. Proteus fuchsinus Pribram, Klassifika- tion der Schizomyceten, Leipzig und Wien, 1933. 73. Serratia indica (Eisenberg) Bergey et al. syn Bacillus indicus ruber Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 285; Bacterium indicum Crookshank, Manual, 1887. 240. Spirillum, parvum Esmarch. (Cent, f. Bakt., I Abt., Orig., 32, 1902. 565; also see Zettnow, ibid., 78, 1916, 1.) From decaying organic matter. Spirillum sputigenum Fliigge. (Lewis, Lancet, Sept. 20, 1884; Fliigge, Die Mi- kroorganismen, 2 Aufl., 1886, 387; Pa- cini a lewi.si Trevisan, I generi e le specie delle Batteriacee, 1889, 24.) From sputum. Staphylococcus activus Pr^vot and Taffanel. (Ann. Inst. Past., 71, 1945, 102.) From puerperal septicemia. An- aerobic. Staphylococcus cilreus duodenalis Gess- ner. (Arch. f. Hyg., 9, 1889, 136.) From the human duodenum. Staphylococcus magnus Black. (Trans. 111. State Dental Soc, 22, 1886, 188.) From the mouth. Staphylococcus medius Black. (Trans. 111. State Dental Soc, 22, 1886, 190.) From the mouth. Staphylococcus pyogenes bovis Lucet. (Ann. Inst. Past., 7, 1893, 327.) From bovine abscesses. Staphylncoccns riscosus Goadby. 702 MANUAL OF DETERMIN^ATIVE BACTERIOLOGY (Mycology of the mouth, London, 1903, 172.) From the mouth. Streptohacterium dextranicum Perquin. (Jour. Microbiol, and Serol., 6, 1940, 226.) Produces slime from sucrose solu- tions. Streptococcus aquatilis Vaughan. (Amer. Jour. Med. Sci., 104, 1892, 184.) From water. Streptococcus liquefaciens Frankland and Frankland. (Phil. Trans. Roy. Soc. London, 178, B, 1888, 264.) From air. After the section covering Strep- tococcus liquefaciens Sternberg emend. Orla-Jensen was in page proof, it was discovered that Frankland and Frank- land had discovered and named a liquefying streptococcus earlier than Sternberg. The Franklands described this species as producing a yellow pigment. Streptococcus pyogenes duodenalis Gessner. (Arch. f. Hyg., 9, 1889, 132.) From the human duodenum. Streptococcus taette (Olsen-Sopp) Bu- chanan and Hammer. (Bacterium lactis longi Troili-Petersson, Ztschr. f. Hyg., 32, 1899, 361 and Milchzeitung, 38, 1899, 438; Streptobacillus taette Olsen-Sopp, Cent. f. Bakt., II Abt., 33, 1912, 9; Buchanan and Hammer, Iowa Sta. Coll. Agr. Exp. Sta., Res. Bull. 22, 1915, 277.) Probably the characteristic organism of Swedish ropy milk. Olsen-Sopp (loc. cit.) misquotes Troili-Petersson's name as Bacillus acidi lactis longus (see Troili-Petersson, Cent. f. Bakt., II Abt., 38, 1913, 1). Thiospira agilissima (Gicklhorn) Ba- vendamm. (Spirillum agilissimum Gickl- horn, Cent. f. Bakt., II Abt., 50, 1920, 418; Bavendamm, Die farblosen und roten Schwefelbakterien, Pflanzenfor- schung, Heft 2, 1924, 116.) From the pond in the Annen Castle Park, Graz, Austria. Contains grains of sulfur. Thiospira elongafa Perfiljev. (Ber. d. Sapropel Komm. Petrograd, 1923, 56.) From mud containing H2S. Thiospira propcra Hama. (Jour. Sci. Hiroshima Univ., Ser. B, Bot., 1, 1933, 157; abst. in Cent. f. Bakt., II Abt., 91, 1934, 200.) Thiospira sulfurica Issatchenko. (Biological observations on the sulfur bacteria (Russian), about 1927, 16 pp.) Vibrio adaptatus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 258.) From sea water and marine sediments. Vibrio agarlyticus Cataldi. (Rev. d. Inst. Bact. (D.N.H.), Buenos Aires, 9, 1940, 375.) From activated sludge. Digests agar. Vibrio albensis Lehmann and Neu- mann syn. Vibrio dunbari Holland, Jour. Bact., 5, 1920, 226; probably Vibrio phosphor escens Jermoljewa, Cent, f. Bakt., I Abt., Orig., 100, 1926, 170; not Vibrio phosphorescens Holland, loc. cit. Vibrio algosus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 257.) Associated with marine kelp. Vibrio arnphibolus Trevisan. (Babes, Ztschr. f. Hyg., 5, 1888, 183; Trevisan, I generi e le specie delle Batteriacee, 1889, 23.) Anaerobe. Vibrio auidus Humm. (Duke Univ. Marine Lab., North Carolina, Bull. 3, 1946, 54.) From intertidal sand, Beau- fort, North Carolina. Digests agar. Vibrio choleroides a and /3 Bujwid syn. Bacterium choleroides Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 131. Vibrio costicolus Smith and Vibrio costicolus var . liquefaciens Smith . (Roy. Soc. Queensland, Proc. for 1937, 49, 1938, 29 and 32.) From tainted ribs of bacon and tank brines in bacon factories. Active growth in 4 to 15 per cent brines. Vibrio euprima, Vibrio yasakii, Coc- cobacillus tolega and Coccobacillus sepiola Majima, Sci-i Kwai- Med. Jour. 50, 1931, 41-67; see Warren, Jour. Bact., 49, 1945, 548.) Phosphorescent bacteria. Vibrio fortis Humm (loc. cit., 55). From seaweed (Gracilaria confervoides) . Digests agar. APPENDIX TO SUBORDER EUBACTERIINEAE 703 Vibrio frequens Humm {loc. cit., 56). From marine algae {Cladophoropsis, Laurencia poitei, etc.) Digests agar. Vibrio halonitrificans Smith. (Ro}^ Soc. Queensland, Proc. for 1937, 49, 1938, 29.) From tank brines in bacon fac- tories. Active growth in 4 to 10 per cent brines. Vibrio haloplanktis ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 261.) Sessile form found associated with marine phyto- plankton. Vibrio hyphalus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography^ Univ. Calif., 5, 1944, 277.) From marine bottom deposits. Vibrio marinagilis ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 264.) From sea water and marine mud. Vibrio marinoflavus ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 258.) From sea water. Vibrio marinofulvus ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 262.) From sea water. Vibrio marinopraesens ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 256.) From sea water. Vibrio marinovulgaris ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 5, 1944, 261.) From sea water. Vibrio notus Humm {loc. cit., 56). From intertidal sand, Atlantic Beach, North Carolina. Digests agar. Vibrio perimastrix Alarie. (Alarie, Thesis, MacDonald Coll. McGill Univ., 1945; see Perlin and Michaelis, Sci., 103, 1946, 673.) Will decompose cellulose only in presence of CO2. Vibrio phytoplanktis ZoBell and Up- ham. (Bull. Scripps Inst, of Ocean- ography, Univ. Calif., 6, 1944, 261.) From sea water and marine phyto- plankton. Vibrio pieris Paillot. (Compt. rend. Soc. Biol., Paris, 94, 1926, 68.) From caterpillars of the cabbage butterfly {Pieris brassicae) which had been para- sitized by larvae of Apanieles glomeratus. Vibrio ponticus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 259.) From sea water. Vibrio rinnpel Lode. (Cent. f. Bakt., I Abt., Orig., 35, 1903, 526; see Ballner, Cent. f. Bakt., II Abt., 19, 1907, 572.) From water. Phosphorescent. Vibrio stanieri Humm {loc. cit., 57). From seaweed {Acanthophora spicifera), Miami, Fla. Digests agar. Vibrio turbidus Humm {loc. cit., 57). From seaweed {Gracilaria confervoides) . Digests agar. Vibrio viridans Miller. (Quoted from Miller, Microorganisms of Human Mouth, Phila., 1890, 85; see Miller, Die Mikroorganismen der Mundhohle, Leip- zig, 1889.) From the mouth. Xanthomonas translucens var. phlei- pratensis Wallin and Reddy. (Phyto- pathology, 35, 1945, 939.) The cause of a bacterial streak disease on timothy grass {Phleum pratense). Xanthomonas vignicola Burkholder. (Phytopath., 34, 1944, 431.) From cow- pea, Vigna sinensis. Yersinia van Loghem (Ann. Past. Inst., 72, 1946, 975), a genus proposed to include Pasteurella pestis and P. pseudo- tuberculosis. Ein neuer fiir Thiere path. Mikroorg. aus dem Sputum eines Pneumoniekran- ken, Bunzel and Federn, Arch. f. Hyg., 19, 1893; 326; Bacillus dubius pneumoniae Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 419; Bacterium sub- pneumonicwin Migula, Syst. d. Bakt., 2, 1900, 376; Bacterium dubium Chester, Man. Determ. Bact., 1901, 142. From the sputum of a pneumonia patient. 704 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY XIII. BACILLACEAE FISCHER.* (Jahrb. f. wiss. Bot., 27, 1895, 139.) Rod-shaped cells, capable of producing spores, either with peritrichous flagella or non-motile ; monotrichous flagellation has been reported but is doubtful. Endospores are cylindrical, ellipsoidal or spherical, and are located in the center of the cell, sub-terminally or terminally. Sporangia do not differ from the vegetative cells except when bulged by spores larger than the cell diameter. Such sporangia are spindle- shaped when spores are central, or wedge- or drumstick-shaped when spores are terminal. Usually Gram-positive. Pigment formation is rare. Aerobic, microaero- philic or anaerobic. Gelatin is frequently liquefied. Sugars are generally fermented, sometimes with the formation of visible gas. Some species are thermophilic, i.e., will grow readily at 55°C. Mostly saprophytes, commonly found in soil. A few are animal, especially insect, parasites or pathogens. Key to the genera of family Bacillaceae. I. Aerobic; catalase positive. Genus I. Bacillus, p. 705. II. Anaerobic or microaerophilic ; catalase not known to be produced. Genus II. Clostridium, p. 763. INTRODUCTION TO THE GENUS BACILLUS. In the fifth edition of the Manual, the late F. D. Chester stated: "It is difficult to offer a rational system of classification for the described forms of the genus Bacillus because of the incompleteness of the data". He prepared a splendid review of the literature but naturally could not supply the data that were missing. He stated further that "The majority of the so-called species in the genus have been imperfectly presented, . . . the net result being that there are comparatively few clearly and definitely described species among the many herein recorded. The development of a better knowledge will be a work of the future". He then discussed the type of work that should be done. A reading of his statement is recommended to anyone contem- plating naming a new species. During the past few years, the writer with the assistance of Francis E. Clark and Ruth E. Gordon has made a study of the genus Bacillus along the lines indicated by Chester. Representative cultures have been obtained from various laboratories, institutions, and private collections. Special mention should be made of the private collection of Prof. J. R. Porter, now at the Iowa State University. It contained about 200 named species and was invaluable for the work. As a result of this study, it ap- pears that many species have been differentiated by such simple characters as mucoid, folded, adherent or rhizoid growth, pigment production, the fermentation of a specific carbohydrate, etc. Others have been grouped because of some special physiological activity such as the decomposition of calcium n-butyrate, xylan, cellulose, etc. Ches- ter rightly considered that these physiological groups had no taxonomic value. Species have been characterized upon a broad basis in the present arrangement on the assumption that one species should not dissociate into another species. Since certain characters are more stable than others, these have been used to establish a * Revised by Mr. Nathan R. Smith, U. S. Bureau of Plant Industry Station, Belts- ville, Maryland (Bacillus), August, 1943, and Prof. R. S. Spray, School of Medicine, West Virginia University, Morgantown, West Virginia {Clostridium) , May, 1942. FAMILY BACILLACEAE 705 species pattern. This has reduced the number of species of the mesophilic members of the genus from many poorly defined organisms to a few well characterized and delimited species. Intermediates occur between related species and have been treated as such. The report on which this arrangement is based has recently been published by Smith, Gordon and Clark (U. S. Dept. Agr. Misc. Pub. Xo. 559, 1946, 112 pp.). Some workers maj' think that the cut in the number of species has been too drastic and that certain organisms listed as varieties, morphotypes, or biotypes should be retained as species. This would not be consistent with the newer knowledge of bacteriology that has been developed during the past two decades. No doubt other species occur in nature that are not included herein. But before jumping to the conclusion that a culture is a new species, closely related organisms as well as the isolate should be studied along the lines given by Chester in the fifth edition of the Manual. The production of indole and the formation of H2S have been omitted from the descriptions because these characters have no taxonomic value. Certain other prop- erties, such as colony form, character of the growth on slants, in litmus milk, etc., have a verj' limited value. They are included for the sake of completeness. Genus I. Bacillus Cohn.* (Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 146 and 175.) From Latin bac- ilhim, a small stick. Synonyms: ? Bacirella Morren, Bull. d. Sci. natur et de Geol., No. 27, 1830, 203; ? Metallacter Perty, Zur Kenntniss kleinster Lebensformen, 1852, 180; ? Bacteridium Davaine, Diet. Encyclop. d. Sci. Med., Ser. I, 8, 1868, 21; Pollendera Trevisan, 1884 (see DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 943); Zopfiella Trevisan, Atti della Accademia Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 93; Cornilia Trevisan, I generi e le specie delle Batteriacee, 1889, 21 ; Urobacillus Miquel, Ann Microg., 1, 1889, 517; Bacterium Migula, Arb. bakt. Inst. Karlsruhe, 1, 1894, 237 (not Bacterium Ehrenberg, Symbolae Physicae seu Icones et Descriptiones Ani- malium, etc., Berlin, 1828, 8); Bactrinum, Bactridium, Bactrillum, Clostrillium, Clostrinium and Paracloster Fischer, Jahresb. f. wissenseh. Bot., 27, 1895, 139; Endo- hacteriuvi Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 103; Astasia Meyer, Flora, S4, 1897, 185; Saccharobacter Beijerinck, Cent. f. Bakt., II Abt., 6, 1900, 200; Fenobacter Beijerinck, ibid.; Aplanobacter E. F. Smith, Bact. in Relation to Plant Dis., 1, 1905, 171; Semiclostridium Maassen, Arb. a. d. k. Gesundheitsamte, Biol. Abt., 5, 1905, 5; Myxobacillus Gonnermann, Ztschr. f. Zuckerind. u. Landwirtsch., 36, 1907, 877; Plennobacterium Gonnermann, ibid.; Serratia Vuillemin, Ann. Mycolog., 11, 1913, 521 (not Serratia Bizio, Polenta porporina, Biblio. Ital., 30, 1823, 288); Schaudinnum, Theciobactrum, Zygostasis, Eisenbergia, Migulanum and Rhagadascia Enderlein, Sitzber. Gesell. Naturf. Freunde, Berlin, 1917,309; C ellulobacillus Simola, Ann. Ac. Sc. Fenn., Ser. A, 3^, No. 1 and 6, 1931 (abst. in Cent. f. Bakt., II Abt., 86, 1932, 89); not C ellulobacillus Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 343; Z yvwbacillus Kluyver and Van Niel, Cent. f. Bakt., II Abt., 9Jf, 1936, 369. Rod-shaped bacteria, sometimes in chains. Sporangia usually not different from the vegetative cells. Catalase present. Aerobic, sometimes showing rough colonies and * Revised by Mr. Nathan R. Smith, U. S. Bur. Plant Industry Station, Beltsville, Maryland, \ugust, 1943. 706 MANUAL OF DETElliMINATIVE BACTP^RIOLOGY forming a pellicle on broth. Usually oxidize carbohydrates or proteins more or less completely, often producing slight acidity, without pronounced accumulation of characteristic products. Soil is the most common habitat. The internationally accepted (Jour. Bact., 33, 1937, 445) type species is Bacillus subtilis Cohn emend. Prazmowski. Key to the species of genus Bacillus. I. Mesophilic (good growth at 30°C), aerobic (sometimes also grow at low concentra- tions of oxygen). A. Spores ellipsoidal to cylindrical, central to terminal, walls thin. Sporangia not distinctly bulged. Gram-positive. 1. Diameter of rods less than 0.9 micron. Cells from glucose or glycerol nutrient agar stain uniformly. a. Growth at pH 6.0. Acetylmethylcarbinol produced. b. Gelatin hydrolyzed (Frazier method). Acid from xylose or arabi- nose with ammoniacal nitrogen. c. Starch hydrolyzed. Nitrites produced from nitrates. 1. Bacillus subtilis. d. Black pigment on carbohydrate media only. la. Bacillus subtilis var. aterrimus. dd. Black pigment on tyrosin media only. lb. Bacillus subtilis var. niger. cc. Starch not hydrolyzed. Nitrites not formed from nitrates. 2. Bacillus pumilus. bb. Gelatin not hydrolyzed. No acid from xylose or arabinose. 3. Bacillus coagulans. aa. No growth at pH 6.0. Acetylmethylcarbinol not formed. b. Casein digested. Urease not formed. 4. Bacillus firmus. bb. Casein not digested. Urease produced. 5. Bacillus lentus. 2. Diameter of rods 0.9 micron or more. Cells from glucose or glycerol nutrient agar appear vacuolated if lightly stained. a. Acid from xylose or arabinose with ammoniacal nitrogen. Acetylmethyl- carbinol not produced. 6. Bacillus megatheriuvi. aa. No acid from xylose or arabinose. Acetylmethylcarbinol produced. b. Saprophytic, sometimes pathogenic but not causing anthrax ; usually motile. 0. Growth on agar not rhizoid. 7. Bacillus cereus. cc. Rhizoid growth on agar; usually non-motile. 7a. Bacillus cereus var. mycoides. bb. Pathogenic. Causative agent of anthrax; non-motile. 8. Bacillus anthracis. B. Spores ellipsoidal, central to terminal, walls thick, remnants of sporangium often adhering. Sporangia distinctly bulged, spindle and racket forms. Gram-variable. 1. Acid and gas from carbohydrates. a. Acetylmethylcarbinol produced. Crystalline dextrins not formed from starch. FAMILY BACILLACEAE 707 9. Bacillus polymyxa. aa. Acetylmethylcarbinol not produced. Crystalline dextrins formed from starch. 10. Bacillus viacerans. 2. No visible gas from carbohydrates. a. Saprophytic. Growth on ordinary media. b. pH of glucose, proteose-peptone broth cultures less than 8.0. Citrates not used as source of carbon, c. Starch hydrolyzed. Acid from sucrose with ammoniacal nitrogen. d. Acid from xylose or arabinose with ammoniacal nitrogen. Acetylmethylcarbinol not formed. 11. Bacillus circulans. dd. No acid from xylose or arabinose. Acetylmethylcarbinol produced. 12. Bacillus alvei. cc. Starch not hydrolyzed. No acid from sucrose. 13. Bacillus laterosporus. bb. pH of glucose, proteose-peptone broth cultures 8.0 or higher. Citrates used as source of carbon. 14. Bacillus brevis. aa. Parasitic. No growth on ordinary media. b. Cause of American foul brood of honey bees. 15. Bacillus larvae. bb. Cause of the milky disease of Japanese beetles {PopilHa japonica ^ewm . ) . Type A. 16. Bacillus popilliae. Type B. 17. Bacillus lentimorbus . C. Spores spherical, central to terminal. Sporangia definitely bulged, drum- sticks and spindles. Carbohydrates not attacked. Gram -variable. 1. Growth on nutrient agar without urea or free ammonia. a. Urease not formed. 18. Bacillus sphaericus. aa. Urease produced. 18a. Bacillus sphaericus var. fusiformis. 2. No growth on nutrient agar without urea or free ammonia. Urease formed. 19. Bacillus pasteurii. II. Thermophilic, optimum temperatures 55°C or above ; slight if any growth at 37°C. Aerobic . * A. Spores ellipsoidal to cylindrical, central to terminal; sporangia not distinctly bulged. 1.. Diameter of rods less than 0.8 micron, a. Gas from carbohydrates. 20. Bacillus thermoamylolyticus. aa. No gas from carbohydrates. * The data on the species of this group are so meager that it is not possible to offer a rational system of classification. Many of the characters used for separating the vari- ous species are probably as variable in this group as they have been found to be in the mesophilic group. Lacking a knowledge of the limits of variability and lacking other pertinent data, the present arrangement is regarded as temporary only. 708 MANUAL OF DETERMIXATIVE BACTERIOLOGY b. Growth below 50°C. c. Nitrites from nitrates, often with liberation of nitrogen gas. 21. Bacillus kaustophilus. 21a. Bacillus pepo. cc. No nitrites from nitrates. 22. Bacillus thennoindifferens. bb. No growth below 50°C. 23. Bacillus thermodiastaticus. 2. Diameter of rods greater than 0.8 micron. a. Growth on nutrient agar. b. Remnants of sporangium adherent. 24. Bacillus cylindricus. bb. Remnants of sporangium not adherent. 25. Bacillus robustus. 25a. Bacillus losanitchii. aa. No growth on nutrient agar. 26. Bacillus calidolaclis. B. Spores ellijjsoidal to cylindrical, central to terminal; sporangia distinctly bulged. 1. Diameter of rods less than 0.9 micron. a. Starch hydrolyzed. b. Nitrites from nitrates, sometimes with liberation of nitrogen gas. 27. Bacillus michaelisii. 27a. Bacillus lobatus. 27b. Bacillus thermononliquefaciens. bb. No nitrites from nitrates. c. Action on cellulose not recorded. 28. Bacillus thermotranslucens. 28a. Bacillus stearothermophilus . 28b. Bacillus aerothermophilus. cc. Cellulose hydrolyzed. 29. Bacillus thennocellulolyticus. aa. Starch not hydrolyzed. b. Nitrites from nitrates, sometimes with gaseous nitrogen, c. Milk unchanged. 30. Bacillus thennoalimentophilus. cc. Milk acid, coagulated. 31. Bacillus thermoliquefaciens. 2. Diameter of rods greater than 0.9 micron. a. Starch hydrolyzed. b. No nitrites from nitrates. 32. Bacillus tostus. C. Spores spherical, central to terminal; sporangium not distinctly bulged. 33. Bacillus viridulus. 1. Bacillus subtilis Cohn emend. Praz- schichte und Fermentwirkung einigen mowski. (Cohn, Beitr. z. Biol. d. Pflan- Bakterien-Arten. Inaug. Diss., Leipzig, zen,;,Heft2, 1872, 174; Heft 3, 1875, 188; 1880.) From Latin subtilis, thin, 2, Heft 2, 1876, 249; Prazmowski, Unter- slender, suchungen iiber die Entwicklungsge- The identity of this species has been FAMILY BACILLACEAE 709 the subject of some controversy owing to the indefiniteness of the original descrip- tions, to the distribution of cultures un- der the name Bacillus siibtilis that were incorrectly identified, to variations in the forms of growth that may be observed, and to confusion with Bacillus cereus. In cases where Bacillus subtilis is said to be "anthrax-like," or "similar to the anthrax bacillus," it should be remem- bered that these terms apply to Bacillus cereus and not to Bacillus subtilis. Conn (Jour. Inf. Dis., 46, 1930, 341) concluded that the so-called Marburg strain fitted the earliest recognizable description of this species which is that given by Prazmowski (loc. cii.), and his view was accepted after a study of cul- tures by the International Committee on Bacteriological Nomenclature (Jour. Bact., 33, 1937, 445). During the past two decades much progress has been made in the study of variations in the stages of growth of bac- teria, the rough, smooth, mucoid, etc., and in the variability in physiology as well. From the recent work of Smith, Gordon, and Clark {loc. cil.) it appears that many species have been character- ized on such simple grounds as growth folded, mucoid, adherent, colored, rhiz- oid, etc., all of which are subject to variation, either induced or spontaneous. The present arrangement of this species is the result of their work combined with data supplied by the work of Conn and others. Species probably identical with or variants of Bacillus subtilis: Bacillus geniculatus de Bary, Beitrag zur Kenntnis der niederen Organismen imMageninhalt, Inaug. Diss., Strassburg, Leipzig, 1885; Bacillus mesentericus fuscus Fliigge, Die Alikroorganismen, 2 Aufl., 1886, 321 {Bacillus ynesentericus Trevisan, I generi e le specie delle Bat- teriacee, 1889, 19; not Bacillus mesenteri- cxis as interpreted by Chester, Del. Agr. Exp. Station 15th Ann. Report, 1903, 86; not Bacillus mesentericus as given by Lawrence and Ford, Jour. Bact., /, 1916, 295);* Bacillus mesentericus vulgatus Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 322 {Bacillus vulgatus Trevisan, I generi e le specie delle Batteriacee, 1889, 19); Bacillus liodermos Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 323 (Ba- cillus No. X, Fliigge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus lactis No. X, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 209 ; Bacillus intermedius Migula, Syst. d. Bakt., 2, 1900, 579; Bacillus cremoris Chester, Man. Determ. Bact., 1901, 274) ; Bacillus laevis Frankland and Frankland, Philos. Trans. Roy. Soc. London, 178, B, 1887, 278 (not Bacillus laevis Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 444); Tyrothrix tenuis Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23 {Bacillus tenuis Trevisan, I generi e le specie delle Batteriacee, 1889, 16); Kartoffelbacillus, Globig, Ztschr. f. Hyg., 3, 1888, 294 {Bacillus roseus Trevisan, loc. cit., 19; Bacillus mesen- tericus ruber Kruse, in Fliigge, Die *T. Gibson, University of Edinburgh (personal communication), has found that the European and supposedly the original strains oi Bacillus mesentericus hydrolyze starch and reduce nitrates to nitrites, whereas the American strains are negative in both of these characters. Furthermore, the latter are usually smooth and when the rough stage exists, it does not resemble a mesentery from which the organism derived its name. This term, however, can still be applied to the European strains. Since the American strains are identical with Bacillus pumilus (Chester, Del. Agr. Exp. Sta- tion, loth Ann. Report, 1903, 87; Lawrence and Ford, Jour. Bact., 1, 1916, 300), it has been recommended (Smith, Gordon, and Clark, loc. cit.) that they be designated as Bacillus pumilus to avoid ambiguity. Since the European Bacillus mesentericus is only a stage of growth of Bacillus subtilis, the former name should be dropped. 710 MANUAL OF DETERMINATIVE BACTERIOLOGY Mikroorganismen, 2, 1896, 199; Bacillus (jlobigii Migula, Syst. der Bakt., 2, 1900, 554; Bacillus vitalis Chester, Man. Determ. Bact., 1901, 286); Bacillus leptosporus Klein, Cent. f. Bakt., 6, 1889, 316; Bacillus No. 6, Pansini, Arch, f. path. Anat. u. Physiol., 122, 1890, 422 (Bacillus coccoideus Migula, Syst. der Bakt., 2, 1900, 558); Bacillus radians Migula, Syst. d. Bakt., 2, 1900, 580 (Bacillus No. IX, Flugge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus lactis No. IX, Kruse, in Fliigge, Die Mikroorgan- ismen, 3 Aufl., 2, 1896, 209; Bacillus stellatus Chester, Man. Determ. Bact., 1901, 274; not Bacillus stellatus Vincent, Ann. Inst. Past., 21, 1907, 69); Bacillus mesentericus pants viscosi II Vogel, Ztschr. f. Hyg., 26, 1897, 404 (Bacillus panis Migula,** Syst. der Bakt., 2, 1900, 576) ; Bacillus armoraciae Burchard, Arb. a. d. bakt. Inst. d. techn. Hochschule zu Karlsruhe, 2, 1898, 46; Bacillus idosus Burchard, ibid., 47; Bacillus subtilis a. Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 635; Bacillus natto Sawamura, Bull. Coll. Agr., Tokyo, 7, 1906, 108; Bacillus mesentericus var. flaous Lau- bach, Jour. Bact., 1, 1916, 497 (Bacillus flavus Bergey et al., Manual, 1st ed., 1923, 286; not Bacillus flavus Fuhrmann, Cent. f. Bakt., II Abt., 19, 1907, 117); Bacillus trujfauti Truffaut and Bezsso- noff, Compt. rend. Acad. Sci., Paris, 175, 1922, 544; Bacillus mesentericus hydrolyticus Hermann and Neuschul, Biochem. Ztschr., 281, 1935, 219. The name Vibrio subtilis Ehrenberg (Infusionsthierchen als vollkommene Or- ganismen, Leipzig, 1838) seems to have given rise to the species name. Spores : 0.6 to 0.9 by 1.0 to 1.5 microns, ellipsoidal to cylindrical, central or para- central. Germination prevailingly equa- torial . Sporangia: Ovoid to cylindrical, only slightly bulged if at all. Rods : 0.7 to 0.8 by 2.0 to 3.0 microns, single or in short chains, rounded ends, stain uniformly. Motile. Gram-posi- tive. The following variations have been observed: Smaller or larger rods, fila- ments, encapsulated cells (the slimy bread organisms ) , few shadow forms, non- motile and Gram -variable. Rods on glucose nutrient agar store small amount of fat. Gelatin stab: Liquefaction. Agar colonies: Usually rough, finely wrinkled, opaque, dull, adherent, slightly spreading, brownish tinge. Variations may be smooth, .soft, thin, translucent, non-adherent, dendroid, coarsely wrin- kled, creamy-white to yellowish to orange. Agar slants : Growth abundant, flat, spreading, usually has a dull mat surface, finely wrinkled, adherent, becoming slightly brownish. Variations may be coarsely wrinkled or folded, non-adher- ent, smooth, thin, translucent, dendroid, creamj'-white to yellow to orange. Some strains show a greenish fluorescence when grown at 45°C on nutrient agar. Broth : Turbid becoming clear with formation of a tough, wrinkled pellicle. Milk: Slowly peptonized, becoming alkaline. ** There has been confusion about the identity of the so-called slimy bread bac- teria. Lehmann and Neumann (Bakt. Diag., 7 Aufl., 2, 1927, 616) stated that they were interrelated and also more or less closely related to Bacillus mesentericus and to Bacillus vulgatus. Laubach (Jour. Bact., 1, 1916, 501) isolated a strain of Bacil- lus panis that lost its capsules on artificial media, although it still remained slimy. From this and the work of Smith, Gordon and Clark (loc. cit.) it is apparent that the slimy bread organisms are mucoid variants of Bacillus subtilis, which may or may not be encapsulated, and motile or non-motile (see also Bacillus subtilis var. viscosus Chester, Del. Agr. Exp. Station, 15th Ann. Report, 1903, 84). FAMILY BACILLACEAE 711 Milk agar plate: Casein hydrolj'zed. Potato : Growth luxuriant, warty or wrinkled to coarsely folded, whitish to pink or yellow, becoming brownish with age. Nitrites formed from nitrates. Starch is hj^drolyzed. Acid with ammoniacal nitrogen from xylose, arabinose, glucose, fructose, galac- tose, mannose, maltose, sucrose, salicin, glycerol, and mannitol. Usually acid from dextrin. Variable reactions on rhamnose, raffinose, and inulin. Usually' no action on lactose. Acetylmethycarbinol produced. Citrates utilized. Optimum temperature 30° to 37°C. Will usually grow from 50° to 56°C. Aerobic, facultative. Source: Original cultures isolated by Cohn from an infusion of lentils (1872), from a boiled infusion of cheese and white beets (1875), and from boiled hay infu- sions (1876). Hence, frequently called the hay bacillus. The folded, non-ad- herent stage of growth (Bacillus vulgatus and the European strain of Bacillus mesentericus) is often called the potato bacillus. Manner of germination of spores established by Prazmowski {loc. cit.). Habitat : Widely distributed in soil and in decomposing organic matter. Note: Bacillus vulgatus has long been separated from Bacillus subtilis by the folded character and the non-adherence of its growth. Recently Lamanna (Jour. Bact., 44, 1942, 611) has attempted to separate this species from Bacillus sub- tilis by the splitting of the spore sheath along the transverse axis upon germina- tion. Since the two species are otherwise morphologicallj" and physiologicall}' alike and since these characters are subject to much variation, there seems to be no valid reason for this separation. One can, if he desires, indicate the different stages of growth; for instance. Bacillus subtilis morphotype vulgatus (or mesentericus) for the folded growth, Bacillus subtilus morphotj'pe panis for the slimy growth, and Bacillus subtilis morphotype globigii for those that produce a red or orange pigment. These terms would apply to the present condition of the culture and would have to be changed if the character of the growth changed. la. Bacillus subtilis var. aterrimus comb. nov. (Potato bacillus, Biel, Cent. f. Bakt., II Abt., 2, 1896, 137; Bacillus aterrimus Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 303; Bacillus viesentericus niger Lunt, Cent. f. Bakt., II Abt., 2, 1896, 572; Bacillus niger Chester, Man. Determ. Bact., 1901, 306.) From Latin aterrimus, very black. Synonyms : Bacillus nigrificans Fabian and Nienhuis, Mich. Agric. Exp. Station, Tech. Bull. 140, 1934, 24; Bacillus tyrosinogenes Rusconi, as referred to by Carbone et al., Inst it. Sierot. Milan., 2, 1921-1922, 29; not Bacillus tyrosinogenes Hall and Finnerud, Proc. Soc. Expl. Biol, and Med., 19, 1921, 48 and Hall. Abstr. Bact., 6, 1922, 6. In the early accounts the production of a blue-black to black pigment on potato was stressed. It was also said to resemble Bacillus subtilis and Bacillus vulgatus on gelatin plates. Recent work (Clark and Smith, Jour. Bact., 37, 1939, 280) has shown that pigmentation occurs only in the presence of a carbohydrate. In addi- tion (Gordon and Smith, Jour Bact., 4^, 1942, 55), it was established that the abil- ity to form the pigment could be lost through serial transfers and colony selection and that the resultant dissoci- ants could not be differentiated from Bacillus subtilis. Source : Isolated from rye bread in moist chamber used for growing some aspergilli (Biel). Habitat: Widely distributed in soil. lb. Bacillus subtilis var. niger comb, nov. {Bacillus lactis niger Gorini, Gior. d. Reale Soc. Ital. Ig., 16, 1894, 9; Bacil- 712 MANUAL OF DETERMINATIVE BACTERIOLOGY his nigcr Migula, Syst. der Bakt., 2, 1900, 636.) From Latin niger, black. The black pigment characterizing this organism is formed only in media contain- ing tyrosine (Clark and Smith, Jour. Bact., 37, 1939, 279). The ability to form the pigment may be lost through serial transfer and colony selection. It then cannot be separated from Bacillus subtilis (Gordon and Smith, loc. cit.). Source: First isolated from milk. Habitat: Widely distributed in soil. 2. Bacillus pumilus Gottheil. (Cent. f. Bakt., II Abt., 7, 1901, 681.) From Latin pumilus, dwarfish, little. Synonyms : Bacillus mesentericus as interpreted by Chester, Del. Agric. Exp. Station, 15th Ann. Report, 1903, 87; Bacillus mesentericus as given by Law- renceand Ford, Jour. Bact., 1, 1916, 295 and 300; Bacillus mesentericus va,r.flavus Laubach, Jour. Bact., 1, 1916, 497; per- haps also Bacillus parvus Neide, Cent. f. Bakt., II Abt., 12, 1904, 344; Bacillus leptodermis Burchard, Arb. a. d. bakt. Inst. d. techn. Hochschule zu Karlsruhe, 2, 1898, 33. Spores: Ellipsoidal to cylindrical, thin walled, naked, central or paracentral, usually about 0.5 by 1.0 micron although some may approach the size of those of Bacillus subtilis. Sporangia: Ellipsoidal to cylindrical, not bulged. Rods : 0.6 to 0.7 by 2.0 to 3.0 microns, usually occurring singly or in pairs. Chains, filaments and shadow forms may be found in some strains . Cells grown on glucose nutrient agar have few small fat globules. Motile with peritrichous fla- gella. Gram-positive. Gelatin stab: Slow liquefaction. Agar colonies: Thin, flat, spreading, dendroid, smooth, translucent. The rough stage also occurs. Agar slants : Growth moderate, smooth, soft, thin, glistening, non-adherent, spreading, usually whitish although it may be yellowish. The rough stage is tough and finely wrinkled, sometimes re- sembling certain strains of Bacillus subtilis. Broth: Lrniform turbidity, with or without a ring or half-formed pellicle. The rough stage forms a pellicle. Milk : Peptonized, sometimes coagu- lated. Milk agar plate: Casein hydrolyzed. Potato : Growth is smooth, thin, spread- ing, moist to slimy, yellowish, turning somewhat brown. The rough stage is dry and finely wrinkled. Nitrites not produced from nitrates. Starch not hydrolyzed. Acid with ammoniacal nitrogen from arabinose, xylose, glucose, fructose, galac- tose, mannose, sucrose, salicin, glycerol and mannitol ; usually also from maltose and raffinose. Reaction variable with dextrin. Usually no acid from rhamnose, lactose, and inulin. Acetylmethylcarbinol produced. Citrates utilized as sole source of carbon. Optimum temperature about 30°C. Maximum temperature allowing growth usually about 50°C. Aerobic . Source: Isolated from plants, cheese, dust, and as a contaminant of media. Habitat : Widely distributed in nature. 3. Bacillus coagulans Hammer. (Iowa Agric. Exp. Station, Research Bull. 19, 1915, 129 ; Sarles and Hammer, Jour. Bact., 23, 1932, 301.) From Latin coagulans, curdling, coagulating. Synonyms: Bacillus thermoacidurans Berry, Jour. Bact., 25, 1933, 72; Bacillus dextrolacticus Andersen and Werkman, Iowa State Coll. Jour, of Sci., 14, 1940, 187. Spores: Ellipsoidal to cylindrical, ter- minal or subterminal, thin walled, 0.6 to 0.9 by 1.0 to 1.5 microns. Sporulation better on acid proteose peptone agar (Stern, Hegarty, and Williams, Food Research, 7, 1942, 186). FAMILY BACILLACEAE 13 Sporangia: Only slightly swollen, if at all. Rods: 0.5 to 0.9 by 2.5 to 3 microns, singly or in short chains, resemble Ba- cillus suhtilis. Cells from glucose agar contain few small fat globules. Motile. Gram-positive. Gelatin: No growth at 20°C. Xo change in gelatin by Frazier method at 45°C. Agar colonies : Small, entire, raised, not characteristic. Agar slants : Growth scant to moderate, thin, flat. On acid proteose peptone agar growth is more abundant and micro- scopically the cells appear healthier. Broth: Moderate uniform turbidity, followed by clearing. Glucose broth attains a pH of 4.0 to 4.4. Milk: Coagulated. Milk agar plate: Weak hjdrolysis of casein. Potato: Growth scant to moderate, thin, spreading, white to cream-colored. May have a sour odor. Nitrites usually not formed from nitrates. Starch is hydrolyzed. Acid from glucose, galactose, fructose, lactose, maltose, sucrose, dextrin, and glycerol. Usually no acid from arabinose and sorbitol. No acid from xylose and mannitol. Organic nitrogen preferable to inorganic. Acetylmethylcarbinol produced. Citrates not used as sole source of carbon. Optimum temperature about 45°C (Hammer, 55°C). Maximum tempera- ture allowing growth 54°C to 60°C. Slow growth, if any, at 25°C. Aerobic, facultative. Source: Isolated from evaporated milk (Hammer) and tomato juice (Berry). Habitat : Canned goods ; probably widelv distributed in nature. Spores: Usually ellipsoidal, central to subterminal, 0.6 to 0.7 by 1.0 to 1.2 microns on Ca-n-butyrate agar (Wer- ner) ; 0.7 to 0.9 by 1.0 to 1.4 microns on nutrient agar. Sporulation better on plain peptone agar than on nutrient agar. Sporangia: Ellipsoidal to cylindrical, sometimes slightly bulged. Rods : 0.6 to 0.9 by 1.5 to 4.0 microns, single or in short chains, few filaments. On glucose nutrient agar there are swol- len, shadow, and other abnormal forms, few small fat globules. Motile with peritrichous flagella. Gram-positive. Gelatin stab : Slow liquefaction. Gela- tin plate shows wide zone of hydrolysis. Agar colonies: Small, smooth, dense, entire, white to pink. Agar slants : Growth moderate, smooth, opaque, not spreading, whitish. Pink variations may occur. Growth inhibited when glucose is added, because of the production of acid. No growth at pH 6.0 or below. Broth: Scant uniform turbiditj' or a flocculent growth. Milk agar plate : Weak to strong casein hydrolysis. Potato: No growth. Nitrites produced from nitrates. Starch is hydroh'zed. Acid from glucose. No acid from arabinose and xylose. Ammonium salts not used as sole source of nitrogen. Acetylmethylcarbinol not produced. Citrates usually not utilized. Urease not produced. Salt tolerance: Will grow in nutrient broth containing 4 to 7 per cent NaCl. Optimum temperature about 28°C. Maximum temperature allowing growth 37°C to 45°C. Source: Seven strains isolated from soils in Central Europe and Egypt. Habitat: Widely distributed in soil. 4. Bacillus firmus Werner. (Cent. f. Bakt., II Abt., 87, 1932, 470.) From Latin firmus, firm, strong. 5. Bacillus lentus Gibson. (Cent. f. Bakt., II Abt., 92, 1935, 368.) From Latin lentus, slow. 714 MANUAL OF DETERMINATIVE BACTERIOLOGY Spores: Ellipsoidal, central to subter- minal, 0.7 to 0.8 by 1.0 to 1.3 microns. Sporangia: Ellipsoidal to cylindrical, may be slightly swollen. Rods : 0.6 1o 0.7 by 2.0 to 3.0 microns, occurring singly or in pairs. Motile with peritrichous flagella. Gram-positive. Gelatin stab : No liquefaction. No change in gelatin by Frazier method. Agar colonies: Small, smooth, entire, glistening, white, opaque. Agar slants : Growth only moderate, slow, thin, gray to white, opaque, not spreading. Nogrowth at pH G.Oor below. Growth inhibited by glucose because of the change to acid reaction. Broth : Faint uniform turliidity, granu- lar sediment. Milk: Unchanged. Milk agar plate: Casein not hydro- lyzed. Potato: No growth. Nitrites not produced from nitrates. Starch is hydrolyzed. Acid from arabinose, xylose, glucose, sucrose, and lactose. Inorganic nitrogen not utilized. Acetylmethylcarbinol not formed. Citrates not used as sole source of carbon. Urease produced . Urea decomposed at room temperature, feebly at 37°C. Salt tolerance : Will grow in nutrient broth containing 4 per cent NaCl. Optimum temperature about 25°C. Maximum temperature allowing growth 37°C. Growth on most media is increased by the addition of urea. Aerobic. Source: Nine strains isolated from soils. Habitat : Common in soils. 6. Bacillus megatherium De Bar\-. {Bacillus 7negaleriuin (sic) Do Bary, Vergleichende Morph. und Biol, der Pilze, 1884, 499.) Generally assumed that the original spelling was a typo- graphical error and that the later spelling megatherium comes from Greek roots meaning big animal (Breed, Science, 70, 1929, 480). Ripple (Arch. Mikrobiol., 11, 1940, 470) holds that the original spelling meaning big rod is the correct form . Synonyms as given by Smith, Gordon, and Clark {loc. cit.) : Bacillus capri Stapp, Cent. f. Bakt., II Abt., 51, 1920, 19; Bacillus carotarum Koch, Bot. Zeit., 18, 1888, 277 (Bacterium carotarum Mig- ula, Syst. d. Bakt., 2, 1900, 293); Ba- cillus cobayac Stapp, Cent. f. Bakt., II Abt., 51, 1920, 10; Bacillus danicus Lohnis and Westermann, Cent. f. Bakt., II Abt., 22, 1908, 258; Bacillus graveolens Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 496 and 535; Bacillus malaharensis Lohnis and Pillai, Cent. f. Bakt., II Abt., 19, 1907, 91; Bacillus musculi Stapp, Cent. f. Bakt., II Abt., 51, 1920, 39; Bacillus oxalaticus Migula, Arb. a. d. bakt. Inst. d. Tech. Hochschule z. Karlsruhe, 1, Heft, 1, 1894, 139; Bacillus petasites Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 535 (Lawrence and Ford, Jour. Bact., 1, 1916, 273); Bacillus ru- minatus Gottheil, ibid., 496; Bacillus silvaticus Neide, Cent. f. Bakt., II Abt., 12, 1904, 32; Bacillus turnescens Zopf, Die Spaltpilze, 1 Aufl., 1883, 66 (Zopfiella turnescens Trevisan, Car. d. ale. nuov. gen. di Batter., 1885, 4). Other possible synonyms given by Neide {loc. cit., 11): Bacterium hirtum Henrici, Arb. bakt. Inst. Karlsruhe, 1, 1894, 44 {Pseudomonas hirtum Ellis, Cent. f. Bakt., II Abt., 11, 1903, 243; Bacillus hirtus Ellis, Ann. Bot., 20, 1906, 233); Bacillus brassicae Pommer, Mitt, botan. Inst. Graz, 1, 1886, 95 {Bacterium brassicae Migula, Syst. d. Bakt., 2, 1900, 296) . Although the name Bacillus turnescens Zopf (which is here regarded as a probable synonym) has priority over Bacillus megatherium, the latter name is preferred because of general usage. Neither of the original descriptions is sufficiently de- tailed to characterize adequately the FAMILY BACILLACEAE 715 species named, and Zopf (Die Spaltpilze, 3 Aufl., 1885, 82-83) regarded the two species as distinct. The modern work on which the present description of Ba- cillus megatherium is based has been largely carried out with cultures identi- fied as Bacillus megatherium, and the true nature of the species is really fixed by the informal emendations made in these more recent descriptions. The emended descriptions give this name a more certain meaning than is given Bacillus tumescens by the descriptions existent in the literature. Spores: Ellipsoidal, sometimes nearly round, central to paracentral, 1.0 to 1.5 by 1.5 to 2.0 microns (larger dimensions have been reported). Sporangia: Ellipsoidal to cylindrical, often in short chains; not swollen. Rods: 1.2 to 1.5 by 2.0 to 4.0 microns, occurring singly and in short chains. Larger and smaller cells, irregular, twisted, and shadow forms are present in some strains, depending upon the sub- strate. Cells from glucose or glycerol nutrient agar usually store much fat and stain unevenly (vacuolated) with dilute stains. Motility with peritrichous fla- gella, usually slow, although some strains may show active motility. Gram-posi- tive. Gelatin stab: Slow liquefaction. Agar colonies : Large, smooth, soft, con- vex, entire, opaque, creamy -white to yellow. The rough stage is usually con- centrically ridged with a thin edge. Agar slants : Growth abundant, soft, butyrous, creamy-white to yellow with pellucid dots. Browning with age ; a few strains become black if the medium con- tains tyrosine. Broth : Medium to heavy uniform turbidity. Milk: Peptonized. Milk agar plate: Casein hydrolyzed. Potato : Growth abundant, smooth, soft to slimy, spreading, creamy-white, pale to lemon-yellow or pink. A few strains are orange-colored, some blacken the potato. The rough stage is wrinkled. Nitrites usually not produced from nitrates. Starch is hydrolyzed. Acid with ammoniacal nitrogen from arabinose, glucose, fructose, sucrose, maltose, de.xtrin, inulin, salicin, glycerol and mannitol. Usually acid from xylose, galactose, mannose, and raffinose; vari- able from lactose. Generally no acid from rhamnose. Acetylmethylcarbinol not formed. Citrates used as sole source of carbon. Uric acid hydrolysis : Variable. Optimum temperature 28°C to 35°C. Maximum temperature allowing growth usually between 40°C and 45°C. Source : Originally isolated from cooked cabbage. Habitat: Widely distributed in soil, water, and decomposing materials. Note : A description of Bacillus mega- therium— Bacillus cereus intermediates follows the description of Bacillus cereus. 7. Bacillus cereus Frankland and Frankland. (Philosoph. Transact. Roy. Soc. London, 178, B, 1887, 279.) From Latin cereus, waxy. Synonj^ms : Bacillus ellenhachensis al- pha Stutzer and Hartleb, Cent. f. Bakt., II Abt., 4, 1898, 31 ; Bacillus ellenhachen- sis Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 540; Bacterium petroselini Bur- chard, Arb. bakt. Inst. Karlsruhe, 2, 1898, 39 (Bacillus petroselini Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 414). The following are given as possible synonyms by Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 540: Bacillus ramosus liquefaciens Flligge, Die Mikroorganis- men, 2 Aufl., 1886, 342; Bacillus stoloni- ferus Pohl, Cent. f. Bakt., 11, 1892, 142 (Bacterium stoloniferus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 91 ; Achromobacter stoloniferum Bergey et al., Manual, 1st ed., 1923, 136); Bacil- lus limosus Russell, Ztschr. f. Hyg., 11, 716 MANUAL OF DETERMINATIVE BACTERIOLOGY 1892, 196 (not Bacillus limosus Klein, Ber. d. deutsch. bot. Gesellsch., 7, 1889, 65; Bacillus limophilus Migula, Syst. d. Bakt., 2, 1900, 550); Bacillus hrevis o Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lutulenius Kern, Arb. bakt . Inst. Karlsruhe, 1, 1897, 402; Bacillus gonio- spoi-us Burchard, Arb. bakt. Inst. Karlsruhe, 3, 1898, 14; Bacterium tur- gescens Burchard, ibid., 18; Bacillus cursor Burchard, ibid., 25; Bacillus lox- osus Burchard, ibid., 37. The following are also listed as syno- nyms or biotypes of Bacillus cereus by Smith, Gordon and Clark: Bacillus ses- silis Klein, Cent. f. Bakt., 6, 1889, 349 and 377 {Bacterium sessile Migula, Syst. d. Bakt., 2, 1900, 290); Bacillus albolactis Migula, ibid., 577 {Bacillus lactis albus Loeffler, Berlin, klin. Wchnschr., 1887, 630); Bacillus lacticola Neide, Cent, f. Bakt., II Abt., 12, 1904, 168 (Bacillus No. V, Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lactis No. V, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 208; Bacterium lacticola Migula, Syst. d. Bakt., 2, 1900, 305; Bacillus excurrens Migula, ibid., 582; Bacillus cereus Chester, Man. Determ. Bact., 1901, 278; not Bacillus cereus Frankland and Frankland, Philos. Trans. Roy. Soc. London, 178, B, 1887, 279) ; Bacillus lactis Neide, Cent. f. Bakt., II Abt., 12, 1904, 337 {Bacillus No. I, Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus fliiggei Chester, Manual Determ. Bact., 1901, 281); Bacillus robur Neide, Cent. f. Bakt., II Abt., 12, 1904, 18; Bacillus thuringiensis Berliner, Zeit. f. angew. Entomol., 2, 1915, 29 (see also Mattes, Gesellschaft zur Beford. der Gesam. Naturw., 62, 1927, 381; Bacterium thuringiensis Chorine, Internat. Corn Borer Invest., 2, 1929, 50); Bacillus cereus var. fluorescens Laubach, Jour. Bact., 1, 1916, 508 {Bacillus fluorescens Bergey et al., Manual, 1st ed., 1923, 298; not Bacillus fluorescens Trevisan, I generi e le specie delle Batteriacee, 1889, 18); Bacillus sublilis Michigan strain. Conn., Jour. Inf. Dis., 46, 1930, 341; Bacillus undulatus den Dooren de Jong, Cent. f. Bakt., I Abt., Orig., 122, 1931, 277 (see also den Dooren de Jong, Arch, f . Mikrobiol., 4, 1933, 36) ; Bacillus siamensis Siribaed, Jour. Inf. Dis., 57, 1935, 143 (see also Bacillus cereus var. siamensis Clark, Jour. Bact., 33, 1937, 435); Bacillus metiens Charlton and Levine, Iowa Eng. Exp. Station, Bull. 132, 1937, 18; (see also Levine, Buchanan and Lease, Iowa State Coll. Jour. Sci., /, 1927, 379); Bacillus tropicus Heaslip, Med. Jour. Australia, 28, 1941, 536. Neide {loc. cit.) gave the following as possible synonyms of Bacillus lacticola: Bacillus butyricus Hueppe, Mitteil. a. d. kaiserl. Gesundheitsamte, 2, 1884, 309; not Bacillus butyricus Macd, Traits de Bact., 1st ed., 1888 {Clostridium butyri- ciim Prazmowski, Untersuchungen iiber die Entwickelungsgeschichte und Fer- mentwirkung einiger Bacterien-Arten. Inaug. Diss., Leipzig, 1880, 23) ; Bacillus aureus Pansini, Arch. f. pathol. Anat. u. Physiol., 122, 1890, 436 (not Bacillus aureus Frankland and Frankland, Philos. Trans. Roy. Soc. London, 178, B, 1887, 272) ; Bacillus lacteus Migula, Syst. d. Bakt., ^, 1900, 571 (No. 17, Lembke, Arch, f. Hyg., 29, 1897, 323); Bacillus gonio- sporus Burchard, loc. cit. Neide also gave the following as possible synonyms of Bacillus lactis: Bacillus lutulentus Kern, loc. cit. ; Bacillus ag- gomeratus Migula, Syst. der Bakt., 2, 1900, 557; Bacillus aniariflcans Migula, ibid., 584; Bacillus cylindrosporus Bur- chard, Arb. bakt. Inst. Karlsruhe, 2 1898,31. Other possible synonyms of Bacillus cereus are: Bacillus anthracoides Hiippe and Wood, Ber. klin. Wchnschr., 16, 1889, 347 (Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 232; Bac- terium anthracoides Migula, Syst. der Bakt., 2, 1900, 281; not Bacterium an- thracoides Trevisan, I generi e le specie delle Batteriacee, 1889, 20); Bacillus pseudanthracis Wahrlich, Bakteriol. Studien, Petersburg, 1890-91, 26 (not FAMILY HAriLLACEAE 717 Bacillus pseudanthracis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 233; Bacterium pseudoanthracis Migula, loc. cit., 282) ; Bacterium flexile Burchard, Inaug. Diss., Karlsruhe, 1897 and Arb. bakt. Inst. Karlsruhe, 3, 1898, 16; Bacil- lus eUenhachi Savvamura, Tokyo Imp. Univ. Coll. Agr. Bull. 7, 1906, 105; Bacillus hoplosternus Paillot, Compt. rend. Acad. Sci., Paris, 163, 1916, 772; Bacillus fulminans Schrire and Green- field, Trans. Roy. Soo. So. Africa, 17, 1929, 309. Spores: Ellipsoidal, average size 1.0 by 1.5 microns (considerable variation has been noted by various writers), central or paracentral, usually freely formed in 24 hours. Germination prevailingly polar. Sporangia: Ellipsoidal or cylindrical, only slightly swollen, if at all. In short to long chains. Rods: 1.0 to 1.2 by 3.0 to 5.0 microns, occurring in long chains, ends square. Cells appear granular or foamy if lightly stained, especially if grown on glucose or glycerol nutrient agar ; fat usually stored. Smooth strains are motile with many peritrichous flagella, rough strains weakh' motile or non-motile. Gram-positive. Gelatin stab: Rapid liquefaction. Agar colonies : Large, flat, entire or irregular, whitish with characteristic appearance by transmitted light de- scribed by various observers as ground glass, moire silk, or galvanized iron. All stages occur from the thin, spreading, very rough and arborescent , to the smooth dense form of colony. Agar slant: Growth abundant, usually non-adherent, spreading, dense, whitish to slightly yellowish. Old slants show characteristic whip-like outgrowths. Some strains produce a yellowish-green fluorescence. Broth: Heavy uniform turbidity, with or without a fragile pellicle. Milk: Rapid peptonization, with or without slight coagulation. Blood serum: Partially liquefied. Hemolysis on blood agar. Potato: Growth abundant, thick, soft, creamy-white to pinkish, spreading over the potato. Rough strains may be folded and more pigmented. Nitrites usually produced from ni- trates. Starch is hydrolyzcd. Acid (with ammoniacal nitrogen) from glucose, fructose, maltose, dextrin, and glycerol. Acid usually from sucrose and salicin. Usually no acid from mannose and lactose. No acid from arabinose, rhamnose, xylose, raffinose, inulin, and mannitol. Acetylmethylcarbinol produced. Citrates usually utilized as sole source of carbon. Optimum temperature about 30°C. Maximum temperature allowing growth varies from 37°C to 48°C, usually about 43°C. Aerobic . Source: From soil, dust, milk, plants, and as contaminant of media. Habitat : Widely distributed. Occurs more often in soil than any other member of the genus. See Chester, Del. Agr. Exp. Station, 15th Ann. Report, 1903, 73; Lawrence and Ford, Jour. Bact., 1, 1916, 284; Conn, N. Y. Exp. Station, Tech. Bull. 58, 1917; Conn and Breed, Jour. Bact., 4, 1919, 273; Soriano, Thesis, LTniv. Buenos Aires, 1935, 569. Bacillus megatherium — Bacillus cereus intermediates. According to Smith, Gordon, and Clark {loc. cit.) intermediate forms occur be- tween Bacillus megatherium and Bacillus cereus which cannot be represented by a distinct species. These intermediates are characterized morphologically by the early appearance on agar of shadow or dis- torted forms, long filaments, and gen- erally only a few spores. Fat globules are smaller and less numerous. Physi- ologically the group is erratic, showing a progression of characters from Bacillus megatherium on the one hand to Bacillus cereus on the other. Acetylmethylcarbi- 718 MANUAL OF DETERMINATIVE BACTERIOLOGY nol and nitrites are not usually formed. Fermentation of the pentoses and manni- tol, the ability to grow well on glucose nitrate agar, susceptibility to the bac- teriophage active against Bacillus megatherium or Bacillus cereus and the general character of the growth deter- mines whether the intermediate is more closely related to Bacillus megatherium or to Bacillus cereus. Bacillus cohaerens Gottheil (Cent. f. Bakt., II Abt., 7, 1901, 458 and 689) may be taken as a representative of this intermediate group resembling Bacillus megatherium more closely than Bacillus cereus. Gottheil gave as possible syn- onyms: Bacillus vermicularis Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 384 (Bacleriutn vermiculare Migula, Syst. d. Bakt., 2, 1900, 302); Bacillus filiformis Tils, Ztschr. f. Hyg., 5, 1890, 293; Bacillus lactis albus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 110; Bacillus virgatus Kern, Arb. bakt. Inst. Karlsruhe, 1, 1897, 416; Bacillus cylindrosporus Burchard, Arb. bakt. Inst. Karlsruhe, 2, 1898, 31; Bacil- lus bipolar is Burchard, ibid., 34. ■ Other strains which apparently belong to this same group are : Bacteriinn pansinii Migula, Syst. d. Bakt., 2, 1900, 303 (Bacillus No. 3, Pansini, Arch. f. path. Anat., 122, 1890, 439; Bacterium granu- latum Chester, Man. Determ. Bact., 1901, 189) ; Bacterium tomentosum Henrici, Arb. bakt. Inst. Karlsruhe, 1, 1897, 40; Bacillus teres Neide, Cent. f. Bakt., II Abt., 12, 1904, 161. Representing those strains in this in- termediate group more closely related to Bacillus cereus is Bacillus simplex Got- theil {loc. cit., 685). Gottheil gave the following as possible synonyms : Bacillus vacuolosis Sternberg, Manual of Bact., 1893, 717; Bacillus natans Kern, Arb. bakt. Inst. Karlsruhe, 1, 1897, 413; Bacillus loxosporus Burchard, Arb. bakt. Inst. Karlsruhe, 2, 1898, 49. 7a. Bacillus cereus var. mycoides (Fliigge) comb. nov. {Bacillus mycoides Flligge, Die Mikroorganismen, 2 Aufl., 1886, 324.) From Greek mykes, fungus; eidos, form, shape, i.e., fungus-like. Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 589, gave the following as probable synonyms: Wurzel bacillus, Eisenberg, Bakt. Diag., 1st ed., 1886, 4; Bacillus figurans Crookshank, Manual, 1st ed., 1886 (Bacterium figurans Chester, Ann. Kept. Del. Col. Agr. Exp. Sta., 9, 1897, 134); Bacillus brassicae Pommer, Mitt, a. d. botan. Inst, zu Graf, 1, 1886, 95 (Bacterium brassicae Migula, Syst. d. Bakt., 2, 1900, 296); Bacterium casei Migula, Syst. d. Bakt., 2, 1900, 304 (Bacillus No. XVI, Adametz, Landw. Jahrb., 18, 1889, 248; Bacterium proteum Chester, Man. Determ. Bact., 1901, 195) ; Bacillus ra7nosus Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 388 (not Bacillus ramosus Veillon and Zuber, Arch. M^d. Exp. et Anat. Path., 10, 1898, 542) ; Bacillus radicosus Zimmer- mann, Die Bakterien unserer Trink- u. Xutzwasser, etc., I Reihe, 1890, 30 (Bacterium radicosum Migula, Syst. d. Bakt., 2, 1900, 283); Bacillus implexus Zimmermann, ibid. (Bacterium im- plexum Migula, ibid., 298); Bacillus in- tricatus Migula, ibid., 546 (Cladothrix in- tricata Russell, Ztschr. f. Hyg., 11, 1892, 191). Another possible synonym is Bacillus praussnitzii Trevisan (I generi e le specie delle Batteriacee, 1889, 20). Laubach, Jour. Bact., 1, 1916, 495, found that this differed from Bacillus mycoides only in the fermentation of lactose. This has been substantiated by later work. Holzmiiller (Cent. f. Bakt., II Abt., 23, 1909, 304) described four varieties of Bacillus mycoides which he designated by Greek letters and in addition named four new species which were apparently only variations of Bacillus mycoides: Bacillus effusus, Bacillus olfactorius, Bacillus nanus and Bacillus dendroides (not Ba- cillus dendr aides Thornton, Ann. Appl. Biol., 9, 1922,247). Bacillus cereus var. mycoides is identi- FAMILY BACILLACEAE 719 cal in all respects with Bacillus cereus except in the following characters : Agar colonies : Grayish, thin, widely spreading by means of long twisted chains of cells, turning to the right or left. Agar slants : Growth thin, rhizoid, grayish, widely spreading, adhering to or growing into the agar. Later, growth becomes thicker and softer. The physiological similarity between Bacillus cereus and Bacillus mycoides has often been noted. Gordon (Jour. Bact., 39, 1940, 98) showed that the rhizoid character of the growth of Bacil- lus mycoides was readily lost by cultiva- tion in flasks containing 100 ml of broth and that the resulting dissociants could not be differentiated from Bacillus cereus. It is, therefore, a question whether Ba- cillus mycoides should be given the dig- nity of a variety of Bacillus cereus or merely designated as a stage of growth (morphotype). Source : Isolated from soil. Habitat: Widely distributed in soil. 8. Bacillus anthracis Cohn emend. Koch. (Les infusories de la maladie charbonneuse, Davaine, Compt. rend. Acad. Sci., Paris, 69, 1864, 393; Cohn, Beitrage z. Biol. d. Pflanzen, 1, Heft 2, 1872, 177; Koch, ibid., 2, Heft 2, 1876, 279; Bacteridie des charbon, Pasteur and Joubert, Compt. rend. Acad. Sci., Paris, S4, 1877, 900; Bacterium anthracis Zopf, Die Spaltpilze, 2 Aufl., 1884, 45; Bacillus {Streptohacter) anthracis Schroeter, Kryptogamen Flora v. Schlesien, 3, 1, 1886, 163; Pollendera anthracis Trevisan, 1884, see Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Bacterium anthracis Migula, in Engler and Prantl, Die natiirlichen Pflanzenfam., 1, la, 1895, 21; Aplanobacter anthracis Erw. Smith, Bacteria in Relation to Plant Diseases, 1905, 171; Bacillus {Bacteri- dium) anthracis Buchanan, Jour. Bact., 3, 1918, 37.) From Greek, gen. of an- thrax, charcoal, a carbuncle, the disease anthrax. According to Smith, Gordon, and Clark {loc. cit.) this species is a pathogenic variety of Bacillus cereus. They worked extensively with the latter but not with many strains of Bacillus anthracis. The only difference between the two seemed to be pathogenicity and motility, and some strains of Bacillus cereus are weakly pathogenic and some practically non- motile. It would appear that Bacilhis cereus is a so-called parent species from which two varieties (var. anthracis and var. mycoides) and several morpho- and biotypes have sprung. Spores : Ellipsoidal, 0.8 to 1.0 by 1.3 to 1.5 microns, central or paracentral, often in chains. Germination polar. Sporangia: Ellipsoidal to cylindrical, not swollen, in chains. Rods : 1.0 to 1.3 by 3 to 10 microns with square or concave ends, occurring in long chains, resemble Bacillus cereus. Cells from glucose or glycerol nutrient agar appear granular (vacuolated) if stained lightly ; many fat globules present. Non- motile. Gram-positive. Gelatin stab : Arborescent in depth, inverted pine tree. Liquefaction cra- teriform becoming stratiform. Agar colonies : Large, irregular, dense, curled structure composed of parallel chains, similar to certain strains of Bacillus cereus. Agar slant : Growth abundant, grayish, dense, spreading, with fimbriate borders. Broth : Little or no turbidity, thick pellicle. Milk: Coagulated, slightly acid, pep- tonized. Potato : Growth abundant, spreading, white to creamy. Nitrites formed from nitrates. Starch is hydrolyzed. Acid from glucose, fructose, sucrose, maltose, trehalose, and dextrin. Some strains produce late and slight acidity in glycerol and salicin. No definite fer- mentation occurs in arabinose, rhamnose, mannose, galactose, lactose, raffinose, inulin, mannitol, dulcitol, sorbitol, inosi- 720 MANUAL OF DETERMINATIVE BACTEKIOLOGY tol, and adonitol (Stein, Vet. Med., 38, 1943). Acetylmethylcarbinol produced. Optimum temperature about 35°C. Maximum temperature allowing growth about 43°C. Aerobic, facultative. Pathogenic for man, cattle, swine, sheep, rabbits, guinea pigs, mice, etc. Source : From blood of infected animals. Habitat. The cause of anthrax in man, cattle, sheep and swine. 9. Bacillus polymyxa (Prazmowski) Migula. {Clostridium polymyxa Praz- mowski, Inaug. Diss., Leipzig, 1880, 37; Migula, Syst. der Bakt., 2, 1900; 638; Granulobacter polymyxa Beijerinck, K. Akad. Wetenschap., Amsterdam, Sec. 2, 1, 1903, No. 10; Granulobacter polymyxa var. mucosu77i and var. tenax Beijerinck and Van Del den. Cent. f. Bakt., II Abt., 9, 1902, 13; further description by Gru- ber, Cent. f. Bakt., II Abt., U, 1905, 353; Aerobacillus polymyxa Donker, loc. cit., 138.) From Greek poly, many or much ; myxa, slime or mucus. This and the species immediately fol- lowing {Bacillus macerans) are sometimes placed in the sub-genus Aerobacillus Donker emend. Kluyver and Van Neil (Donker, Inaug. Diss., Delft, 1926, 138; Kluyver and Van Neil, Cent. f. Bakt., II Abt., 94, 1936, 402; not Aerobacillus Pribram, Jour. Bact., 18, 1929, 374; not Aerobacillus Janke, Cent. f. Bakt., II Abt., SO, 1930,481). For a study of this group and a review of the literature see Porter, McCleskey and Levine, Jour. Bact., 33, 1937, 163. They give the following as synonyms of Bacillus polymyxa: Bacillus asterosporus Migula {Astasia asterospora Meyer, Flora, Erg. Bd., 84, 1897, 185; Migula, Syst. der Bakt., 2, 1900, 528; Aerobacillus asterospoj-us Donker, loc. cit., 141); Bacillus ovoaethylicus Pribram {Bacillus mycoides var. ovoaethylicus Wagner, Ztschr. f. Untersuch. d. Nahrungs- u. Genussmittel, 31, 1916, 234; Pribram, Klassifikation der Schizomyceten, Leip- zig und Wien, 1933, 86); Bacillus aero- sporus Greer, Jour. Inf. Dis., 4^, 1928, 508. Gottheil (Cent. f. Bakt., II Abt., 7, 1901, 727) regarded the following as syno- nyms : Bacillus thalassophilus Russell, Ztschr. f. Hyg., 11, 1892, 190; Bacillus subanaerobius Migula, Syst. der Bakt., 2, 1900, 600. Bredemann (Cent. f. Bakt., II Abt., 23, 1909, 45) admitted that theorganisms, Bacillus asterosporus alpha. Bacillus dilaboides, and Bacillus clostridioides , named by Haselhoff and himself in an earlier article (Landwirtsch. Jahrb., 35, 1906, 420, 426, 432) were merely variants of Bacillus asterosporus. The following is usually considered a variety or strain of Bacillus polymyxa differing from the latter mainly in the production of a violet pigment on potato and agar in the presence of peptone : Bacillus violarius acetonicus Brdaudat, Ann. Inst. Pasteur, 20, 1906, 874 {Aero- bacillus violarius Donker, Inaug. Diss., Delft, 1926, 141). Also a probable synonym of Bacillus polymyxa is Bacillus amaracrylus Viose- net (Bacille de I'amertume, Voisenet, Compt. rend. Acad. Sci., Paris, 153, 1911, 363; Voisenet, Ann. Inst. Pasteur, 32, 1918, 477; Aerobacillus am,aracrylus Donker, loc. cit., 141). The chief charac- ter in which it differs from Bacillus polymyxa is its ability to dehydrate glycerol with the formation of acrolein. Also a probable variant of Bacillus polymyxa is Bacillus pandora Corbet (Jour. Bact., 19, 1930, 321). The chief characters in which the latter differs from the former are the production of acid without gas from glucose and the lack of diastatic action. Spores : Ellipsoidal, 1.0 to 1.5 by 1.5 to 2.5 microns, central to subterminal, wall usually thick and stainable. Freely formed. Sporangia: Swollen, spindle-shaped (Clostridia), sometimes clavate. Rods : 0.6 to 1.0 by 2.5 to 6.0 microns, FAMILY BACILLACEAE 721 occurring singly or in short chains. Cells contain small fat globules when grown on glucose nutrient agar. Motile with peri- trichous flagella. Gram-variable. Gelatin stab: Slow liquefaction. Hj-- drolysis of gelatin always positive bj- Frazier technic. Agar colonies: Thin, inconspicuous, lobed, spreading over entire plate. Rough forms are round, whitish, and sometimes tough. Agar slant : Growth scant to moderate, indistinct to whitish. On glucose agar, the growth is much heavier, raised, gummy, glistening; gas is formed. Broth: Uniform to granular turbidity, flocculent to slimy sediment. Rough stage forms pellicle. Final pH of glucose broth cultures 5.2 to 6.8. Milk: Not coagulated, gas usually formed . Milk agar plate: Casein hydrolyzed. Potato : Growth moderate to abundant, whitish to light tan, potato decomposed with formation of gas. Growth of rough stains is denser and heaped up. Nitrites are produced from nitrates. Starch is hydrolyzed. Crystalline de.\- trins are not produced. Acid and gas (with ammoniacal nitro- gen) from arabinose, xylose, glucose, fructose, galactose, mannose,' maltose, sucrose, lactose, trehalose, cellobiose, raffinose, melezitose, de.xtrin, inulin, salicin, glycerol, and mannitol. Gum is also usually formed. Erythritol, adoni- tol, dulcitol and inositol not fermented. With organic nitrogen no acid or gas from rhamnose or sorbitol (Porter, IMcCleskey, and Levine, loc. cit., also Tilden and Hudson, Jour. Bact., J^, 1942, 530). This, however, could not be confirmed by Smith, Gordon, and Clark {loc. cit.) who found that acid and gas were produced from both carbohydrates. Hemicellulose and pectin are attacked (Ankersmit, Cent. f. Bakt., I Abt., Orig., JtO, 1905, 100). In glucose broth, ethyl alcohol and butylene-glycol are produced also small amounts of acetone and butyl alcohol. Acetylmethylcarbinol is produced. Citrates usualh' not utilized as sole source of carbon. Optimum temperature about 30°C. No growth at 42°C to 45°C; good growth at 20°C, slow at 13°C. Not agglutinated by Bacillus inacerans sera, results with homologous sera irregu- lar (Porter, McCIeskej^ and Levine, loc. cit.). Aerobic, facultative. Source : First isolations were from grain, soil, and pasteurized milk. Habitat : Widely distributed in water, soil, milk, feces, decaying vegetables, etc. In addition see: Chester, Del. Agr. Exp. Station, 15th Ann. Report, 1903, 65; Wund, Cent. f. Bakt., I Abt., Orig., 4-2, 1906, 193, 289, 385; Wahl, Cent. f. Bakt., II Abt., 16, 1906, 489; Ritter, Cent. f. Bakt., II Abt., 20, 1908, 21 ; Meyer, Cent, f. Bakt., I Abt., Orig., Jf9, 1909, 305; Bredemann, Cent. f. Bakt., II Abt., 23, 1909, 41 ; Virtanen and Kurstom, Bio- chem. Ztschr., 161, 1925, 9; Stapp and Zycha, Arch. f. Mikrobiol., 2, 1931, 493; Zycha, Arch. f. Mikrobiol., 3, 1932, 194; Patrick, Iowa State Coll. Jour. Sci., 7, 1933, 407. 10. Bacillus macerans Schardinger. (Rt)ttebazillus 1, Schardinger, Wiener klin. Wochenschr., 17, 1904, 207; Schard- inger, Cent. f. Bakt., II Abt., H, 1905, 772; Aerobacillus macerans Don- ker, Inaug. Diss., Delft, 1926, 139; Zymo- bacillus macerans Kluyver and Van Niel, Cent. f. Bakt., II Abt., 94, 1936, 402.) From Latin macerans, softening, macer- ating or retting. Porter, McCleskey, and Levine, Jour. Bact., 33, 1937, 163, regard the following as a synonym of Bacillus macerans: Bacillus acetoethylicum Northrup, Ashe, and Senior, Jour. Biol. Chem., 39, 1919, 1 {Aerobacillus acetoethylicus Donker, loc. cit.). The following is probably a variant of 722 MAXUAL OF DETERMINATIVE BACTERIOLOGY Bacillus macerans: Aerohacillvs sclunjl- killiensis Eisenberg, Jour. Amer. Water Works Assoc, S4, 1942, 365. It is said to differ from Bacillus macerans in that sorbitol is not fermented, hydrogen sul- fide is produced and gelatin is liquefied. Spores: Ellipsoidal, 1.0 to 1.5 by 1.5 to 2.5 microns, terminal to subterminal ; wall thick and stainable. Sporangia : Swollen terminally, clavate. Rods : O.G to 1.0 by 2.5 to 6.0 microns, occurring singly or in pairs , cells are larger on sugar media than on sugar-free media, and contain a few small fat globules. Motile. Gram-variable. Gelatin stab: Liquefaction variable (see optimum temperature). Gelatin is hydrolyzed as determined by the Frazier technic (30°C). Agar colonies : Small, thin, transparent to whitish, irregular, usually smooth. Agar slant : Growth moderate, spread- ing, inconspicuous. Broth: Turbid, slight sediment. In sugar broths some strains produce slime. Glucose broth cultures, pH 5.0 to 5.5. Milk: Acid and gas. No visible peptonization. Milk agar plate: Casein not hydro- lyzed in one week; later usually slight hydrolysis. Potato : Growth indistinct, gas is formed and the potato is digested. Fruity odor sometimes produced. Nitrites produced from nitrates. Starch is hydrolyzed. Acid and gas from arabinose, rhamnosc, xylose, glucose, fructose, galactose, man- nose, sucrose, maltose, lactose, trehalose, cellobiose, raffinose, melezitose, dextrin, inulin, salicin, pectin, xylan, glycerol, mannitol, and sorbitol. Erythritol, adonitol, dulcitol, and inositol not fer- mented (Porter, McCleskey, and Levine, loc. cit.). Produces acetone and ethyl but never butyl alcohol ; ratio acetone to alcohol is 1:2. Acetylmethylcarbinol not produced. Citrates not utilized as sole source of carbon. Optimum temperature about 37°C. Good growth at 42° to 45°C and some- times slightly higher ; poor growth, if any, at20°C. Differentiated from Bacillus polymyxa by the production of crystalline dextrins from starch, lack of formation of acetyl- methylcarbinol, and by growth at 42°C to 45°C. All strains agglutinated by homologous sera but not by Bacillus pohjmyxa serum. Aerobic, facultative. For additional literature, see Porter, McCleskey and Levine, Jour. Bact., 33, 1937, 180. Source : Originally isolated from vats in which flax was retting. Habitat: Widely distributed in soil, water, decomposing starchy materials, retting flax, etc. 11. Bacillus circulans Jordan. (Jor- dan, Exp. Inv., Mass. State Board Health, Part II, 1890, 831; Bacterium circulans Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 92; also see Ford, Jour. Bact., 1, 1916, 519.) From Latin circulans, making round or circular. Smith, Gordon, and Clark (loc. cit.) consider Bacillus circulans as a complex (see also Gibson and Topping, Soc. Agric. Bact. (British), Abstr. Proc, 1938, 43) because of the variations in the character of the growth and quantitative differences in physiology. All stages of growth may be found from the smooth actively motile strains that have motile colonies to the mucoid, non-spreading strains. The spe- cies is more saccharolytic than proteo- lytic, considerable variation being found in its action on gelatin and casein. The following are regarded as variants : Bacil- lus closteroides Gray and Thornton, Cent. f.Bakt.,IIAbt., 73, 1928, 93; Bewegungs- typus schwarmender Bakterien, Russ- Munger, Cent. f. Bakt., I Abt., Orig., 142, 1938, 175; Bacillus krzemieniewski FAMILY BACILLACEAE 723 Kleczkowska, Norman and Snieszko, Soil Sci., 4-9, 1940, 185 (a mucoid variant ) . Also probable variants : Bacillus platus Soriano, Thesis, Univ. Buenos Aires, 1935, 572; Bacillus navifonnis Soriano, ibid., 574 (not Bacillus navifonnis Jungano, Comp. rend. Soc. Biol., Paris, 1, 1909, 122). Spores : Ellipsoidal, 0.8 to 1.2 by 1.1 to 2.0 microns, terminal or subterminal. Spore wall thick and stainable. In some strains spores may be kidney -shaped and remnants of sporangium may adhere. Sporangia : Swollen terminally, clavate. Rods : 0.5 to 0.9 by 2.0 to 5.0 microns, sometimes curved, usually occurring singly. Cells contain small fat globules when grown on glucose agar. Motile, some strains exceedingly so. Gram- variable, usually negative. Gelatin stab : Slow cone-shaped lique- faction, liciuefied portion evaporating (Jordan) ; no liquefaction (Ford). Gela- tin hydrolyzed if tested by the Frazier method. Agar colonies: Thin, transparent, spreading over entire surface of plate. Often nearly invisible. The colonies of the rougher or mucoid strains are small, entire, whitish, non-spreading. Giant agar colonies : If the surfaces of agar plates are dried before inoculation with very motile strains, instead of spreading as a thin layer of individual cells, minute rotating colonies proceed out from the edge of the colony, some- times becoming entirely disconnected from it. In moving out across the agar surface, non-motile cells are left behind. These vcv&y grow later. Eventually the whole plate is covered. Agar slant: Growth thin, transparent, spreading, becoming denser. Mucoid strains are dense, non-spreading, entire, gummy and adherent. Broth: Light to fair turbidity with flocculent to slimy sediment. Some strains do not grow perceptibly. In glu- cose broth cultures the final pH is usually 5.0 to 5.8. Milk: Usually acid, slowly coagulated. Milk agar plate: Casein not hydro- lyzed. Weak hydrolysis with some strains. Potato: Growth is very variable, from none to good growth, from colorless to yellowish, pink, or brownish. Nitrites usually produced from nitrates . Starch is hydrolyzed. Crystalline dex- trins usuall}' not formed. Acid without gas (with ammoniacal nitrogen) from glucose, fructose, man- nose, galactose, sucrose, maltose, raffi- nose, salicin, and dextrin. Usually acid from arabinose, xylose, lactose, glycerol, and mannitol. Reaction variable with rhamnose and inulin. Acetylmethylcarbinol not produced. Citrates usually not utilized. Methylene blue reduced and then reoxidized. Urease produced by some strains. Optimum temperature about 30°C. Maximum temperature allowing growth, 40°C to 48°C. A few strains will grow up to 52°C. This species is closely related to Bacil- lus macerans from which it is distin- guished by the lack of gas formation from carbohydrates and the lack of crystalline dextrins from starch. It is also close to Bacillus alrei as indicated by the key. Source : Found occasionally in tap water, Lawrence, Mass. (Jordan). Habitat: Widely distributed in soil, water, and dust. 12. Bacillus alvei Cheshire and Chej'ne. (Jour. Roy. Mic. Soc, Ser. II, 5, 1885, 592.) From Latin alveus, bee- hive. Probabl}^ identical with the above : Bacillus paraalvei Burnside, Amer. Bee Jour., 72, 1932, 433; Burnside and Foster, Jour. Econ. Entomol., 28, 1935, 578. Spores : Ellipsoidal, 0.7 to 1.0 by 1.5 to 2.5 microns, central to terminal. Free spores freciuently lie in parallel arrange- ment like the rods. 724 MANUAL OF DETERMINATIVE BACTERIOLOGY Sporangia: Swollen, spindle-shaped to clavate. Rods: 0.5 to 0.8 by 2 to 5.0 microns. Cells frequently lie parallel, side by side, like cartridges in a clip. Usually non- capsulated and very motile. Few small fat globules in cells from glucose agar. Gram-variable (young cells Gram-posi- tive, becoming Gram-negative). Gelatin stab: Slow liquefaction. Agar colonies : Thin, translucent, smooth, quickly spreading as a thin layer over entire plate. The growth thickens with age. Rough and mucoid strains do not spread. Giant agar colonies : If the surfaces of agar plates are dried before inoculation with the motile strains, instead of spread- ing as a thin layer, minute bullet-shaped colonies proceed out from the edge of the colony and move across the sterile agar. Non-motile and sometimes motile cells are left behind along the path made by the motile colony (Smith and Clark, Jour. Bact., S5, 1938, 59). Eventually the whole plate is covered. Agar slant : Growth thin, inconspicu- ous, spreading, becoming thicker. Rough and mucoid strains do not spread, growth is heaped, and sometimes gummy. Broth: Uniform turbidity. Rough strains may form a pellicle. Glucose broth cultures have a pH of 5.0 to 6.0. Milk: Usually coagulated, little or no .acid, peptonized. Milk agar plate: Casein hydrolyzed. Potato : Growth scant to moderate, soft, spreading, usually creamy yellow. Nitrites not produced from nitrates. Starch is hydrolyzed. Acid (with ammoniacal nitrogen) from glucose, fructose, galactose, sucrose, maltose, de.Ktrin and glycerol. Reaction variable on mannose, lactose, rafhnose, salicin, and mannitol. No acid from arabinose, rhamnose, xylose, and inulin. Acetylmethylcarbinol is produced. Citrates not utilized. Optimum temperature about 30°C. Maximum temperature allowing growth 43°C to 45°C. Putrefactive odor on media rich in proteins (egg). Aerobic. Source : Isolated from diseased brood of bees. Habitat : Associated with European foulbrood of honey bees; widely distrib- uted in soil. Note: The following must be con- sidered in connection with Bacillus alvei: Bacilhis phiion White. (U. S. Dept. of Agric, Bur. Entomol., Circ. 157, 1912; Diplococcus pluion Bergey et al.. Manual, 2nd ed., 1925, 45.) See also Lochhead, Science, 67, 1928, 159 andProc. IV Intern. Congr. Entomol., 2, 1929, 1005; Burnside, Jour. Econ. Entomol., 27, 1934, 656; Tarr, Ann. Appl. Biol., 24, 1935, 614; Burri, Beihefte z. schweiz. Beinenz., 1, Heft 1, 1941. White considered Bacillus pluton to be the cause of European foulbrood though the evidence was indirect since the organ- ism was not cultivated. Lochhead sug- gested that Bacillus pluton and Strepto- coccus apis are variants or stages in the life history of Bacillus alvei, a hypothesis supported by Burnside who included, in addition. Bacterium eurydice. Accord- ing to Burri, rod forms identical with Bacterium eurydice give rise to Bacillus pluton which is not directly cultivable. Tarr considers European foulbrood to be caused by Bacillus pluton, distinct from Bacillus alvei, and considers it a strict parasite able to multiply only in the intestines of young larvae. Source : Larvae of the honey bee in- fected with European foulbrood. 13. Bacillus laterosporus Laubach. (Jour. Bact., 1, 1916, 511.) From Latin latus, latcris, the side; Greek sporus, seed; M.L., spore. Synonym: Bacillus orpheus White. (U. S. Dept. of Agric, Bur. Entomol., Circ. 157, 1912, 3.) Although named by White, the organism was not described FA\nLY BACILLACEAE 725 until 1917 (McCray, Jour. Agr. Research, 8, 1917, 410). Resembles Bacillus latero- sporus (White, U. S. Dept. Agric. Bull. 810, 1920, 14). According to the rules of priority, the name to be used is Bacillus laterosporus. Spores : Ellipsoidal, 1.0 to 1.3 by 1.2 to 1.5 microns, central to subterminal, formed close to one side, remnants of the sporangium adhering to the other side. Sporangia: Swollen, spindle-shaped. Rods : 0.5 to 0.8 by 2.0 to 5.0 microns, occurring singly and in pairs. Ends pointed or poorly rounded. Cells from glucose nutrient agar may have few small fat globules. Motile. Gram-variable. Gelatin stab: Slow liquefaction. Agar colonies : Thin, transparent, irreg- ular, spreading. Colonies of rough strains are small, round, convex, non- spreading. Agar slant: Growth moderate, flat, translucent to opaque, moist, sometimes with a silvery sheen. Broth : Uniform to granular turbidity ; usually no pellicle. Glucose broth cul- tures, pH 6.0 to 6.4. Milk: Usually curdled, peptonized. Milk agar plate: Occasionally weak hydrolysis of casein. Potato: Growth thin, spreading, gray- ish to pinkish, turning light brown with age. Sometimes finely wrinkled. Nitrites produced from nitrates. Starch is not hydrolyzed. Acid (with ammoniacal nitrogen) from glucose, fructose, maltose, glycerol, and mannitol. Reaction variable on galac- tose, mannose, and salicin. No acid from arabinose, rhamnose, xylose, sucrose, lactose, raffinose, inulin, and dextrin. Acetylmethylcarbinol not produced. Citrates not utilized as sole source of carbon. Optimum temperature about 28°C. Maximum temperature allowing growth 37°C to 45°C. Aerobic. Source : Isolated from water. Habitat: Widely distributed in soil, water and dust. 14. Bacillus brevis Migula emend. Ford. (Bacillus No. I, Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus laciis No. I, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 208; Migula, Syst. d. Bakt., 2, 1900, 583; not Bacillus brevis Frankland and Frankland, Microorgan- isms in Water, 1894, 429; Bacillus fliiggei Chester, Man. Determ. Bact., 1901, 281; Ford, Jour. Bact., /, 1916, 522.) From Latin brevis, short. Synonym : Bacillus centrosporus Ford, Jour. Bact., /, 1916,524. There is some doubt as to the identity of Migula's Bacillus brevis which origi- nally was Fliigge's Bacillus No. I. Neide (Cent. f. Bakt., II Abt., 12, 1904, 337) also renamed Fliigge's organism. He called it Bacillus lactis and described it sufficiently that it may be recognized as a strain of Bacillus cereus. Ford believed that his isolations from milk, soil and dust conformed to Migula's description of Bacillus brevis. Ford's interpretation has been accepted. The species has ap- parently become well established in Europe (Gibson and Topping, Soc. Agric. Bact. (British), Abstr. Proc, 1938, 43) as well as in America. Description from Ford and from Smith, Gordon, and Clark {loc. cit.). Spores : Ellipsoidal, 1.0 to 1.3 by 1.5 to 2.0 microns, central to subterminal. Spore walls thick and stainable. An occasional strain shows the relationship of this species to Bacillus laterosporus in that some of the spores may be lateral and remnants of the sporangia may adhere to one side of the spore. Sporangia: Definiteh' swollen, spindle- shaped to clavate. Rods: 0.4 to 0.8 by 1.5 to 5.0 microns, with pointed ends, occurring singly or in pairs. On glucose agar cells contain numerous small fat globules. Motile. Gram-variable. Gelatin stab: Slow liquefaction. Agar colonies : Thin, flat, translucent, smooth, quickly spreading over plate. Agar slants : Growth smooth, moist, spreading, grayish white. 726 MANUAL OF DETERMINATIVE BACTERIOLOGY Broth: Usually heavy uniform tur- bidity, sometimes with a fragile pellicle. Glucose broth cultures have a pH of 8.0 to 8.6 after 7 days. Milk: Peptonized. Milk agar plate: Casein hydrolyzed. Potato : Growth scant to moderate, flat, spreading, soft, creamy-yellow to pink to brownish. Nitrites usually formed from nitrates. Starch is not hydrolyzed. Acid (with ammoniacal nitrogen) from glucose, fructose, maltose, and sucrose. Usually acid from galactose and glycerol. Reaction is variable on rhamnose and mannitol. No acid from arabinose, xylose, mannose, lactose, raffinose, inulin, dextrin and salicin. With organic nitrogen, the acid formed from carbohydrates is masked bj^ the alkalinity due to proteolysis. Acetylmethylcarbinol is not produced. Citrates usually utilized as a sole source of carbon. Optimum temperature about 30°C. Maximum temperature allowing growth varies from 43°C to 54°C. Produces antibiotic substances (tyro- thricin, gramicidin ; see Dubos and Hotchkiss, Jour. Exp. Med., 73, 1941, 629). Aerobic. Source: From milk (Fliigge); from milk, soil and dust (Ford). Habitat: Widelj^ distributed in .soil, water, dust, milk, etc. 15. Bacillus larvae White. (Bac- terium X, Moore and White, New York State Dept. Agr., 11th Ann. Rept. Com. Agr. for 1903, 1904, 111; Bacillus X, Moore and White, ibid., Rept. for 1904, 1905, 106; White, Thesis, Cornell Univ., Ithaca, N. Y., 1905; White, U. S. Dept. Agr. Bur. Entomol., Tech. Ser. Bui. 14, 1906, 32; White, U. S. Dept. Agr. Bui. 809, 1920, 13.) From Latin larva, a ghost; M. L., of a larva. Synonyms : Bacillus hrandenhurgiensis Maassen,Mittl.a. d. kaiserl. biol. Anstalt f. Land- u. Forstw. in Dahlem, Heft 2, 1906, 28; ibid., Heft 7, 1908, 24 pp.; Arb, a. d. Anstalt f. Land- u. Forstw., 6, 1908. 61; Bacillus burri Cowan, British Bee- keeper's Guide Book. 20th ed., London, 1911, 171. Description from Lochhead, Sci. Agr., .9, 1928, 84. Spores: Ellipsoidal, central to sub- terminal. Sporangia: Swollen, spindle form. Rods : 0.5 to 0.8 by 2.5 to 5.0 microns, occurring singly and in chains. Motile. Gram-variable. Gelatin stab : No growth. In carrot- gelatin, slow liquefaction. Yeast-carrot agar colonies : Small , whit- ish, somewhat transparent, smooth, slightly glistening. Agar slant : No growth. With addition of carrot extract, noticeable growth along line of inoculation. More abundant growth if yeast extract is also added. Yeast-carrot broth : Fungoid in appear- ance, floating masses which may be broken up by shaking to produce a uni- form clouding. Carrot-milk: Acid with curdling. No peptonization . Potato : No growth. Nitrites formed from nitrates (see Lochhead, Can. Jour. Res., C, 15, 1937, 79). Starch not hj^drolyzed (carrot-starch agar). Acid (in yeast extract-peptone broth) from xylose, glucose, fructose, galactose, salicin. Slight acidity by some strains from lactose and sucrose. No acid from mannitol or dulcitol. Thiamin replaces the growth factor in vegetable or yeast extracts, etc. (Loch- head, Jour. Bact., U, 1942, 185). Optimum temperature about 37°C. Maximum temperature about 45°C. Source : From diseased brood. Habitat : Causal organism of American foulbrood of honey bees. See in addition : White, Science, 4^' 1919, 362; Sturtevant, Jour. Agr. Res.,.?S, 1924, 129; Borchert, Die seuchenhaften Krankheiten der Honigbiene. Berlin, FAMILY BACILLACEAE 727 1926; Sturtevant, Jour. Agr. Res., 4^, 1932, 257; Hitchcock, Jour. Econ. Ento- mol., 29, 1936, 895; Stoilowa, Cent. f. Bakt., II Abt., 99, 1938, 124; Tarr, Ann. Appl. Biol., 25, 1938, 633; Hoist and Sturtevant, Jour. Bact., 40, 1940, 723. 16. Bacillus popilliae Dutky. (Jour. Agr. Research, 61, 1940, 59.) From the genus name of the Japanese beetle, Popil- lia japonica Newm. Spores : Cylindrical, 0.9 by 1.8 microns, central. Free spores have not been observed. Sporangia: Swollen, spindle-shaped. Contains a refractile body at the broader pole of the cell which is about half the size of the spore and reacts similarly to stains. Rods : Unstained, 0.9 by 5.2 microns. Stained by crystal violet after fixing in Schaudinn's solution, 0.3 by 3.5 microns. Non-motile. Gram-positive. Unheated egg yolk-beef infusion agar slants : Growth occurs as small discrete colonies. Optimum temperature about 30°C. Maximum temperature about 36°C. Aerobic, facultative. Source : From infected larvae. Habitat : Cause of type A milky disease of Japanese beetle, Popillia japonica. See Hawley and White, X. Y. Ent. Soc. Jour., 43, 1935, 405. 17. Bacillus lentimorbus Dutky. (Jour. Agr. Res., 61, 1940, 65.) From Latin lentus, slow, lingering, and morbus, death. Spores : Ellipsoidal, 0.9 by 1.8 microns, central. Sporangia: Swollen, spindle-shaped. No refractile granule at pole. Rods : Unstained, 1.0 by 5.0 microns. Stained by crystal violet after fixing in Schaudinn's solution, 0.5 by 4.0 microns. No growth on artificial media. Optimum temperature about 25°C. Maximum temperature about 30°C. Aerobic, facultative. Source : Diseased larvae. Habitat : Cause of type B milky disease of Japanese beetle, Popillia japonica. 18 . Bacillus sphaericus Neide. (Cent . f. Bakt., II Abt., 12, 1904, 350.) From Latin sphaericus, spherical. Neide {loc. cit.) gave the following as possible synonyms : Bacillus (Strepto- bacter) albuminus Schroeter, in Cohn, Kryptogamenflora von Schlesien, 3, 1, 1886, 162; Bacillus putrificus coli Fliigge, Die Mikroorganismen, 2 Aufi., 1886, 303; Bacillus gracilis Zimmermann, Die Bak- terien unserer Trink- u. Nutzwasser, etc., 1, 1890, 50 {Bacterium gracile Ches- ter, Man. Determ. Bact., 1901, 198); Bacillus butyricus Botkin, Ztschr. f. Hyg., 11, 1892, 421; Bacillus thalas- sophilus Russell, Ztschr. f. Hyg., 11, 1892, 190; Bacillus pseudotelanicus aero- bius Kruse, in Fliigge, Die Mikoorganis- men, 3 Aufl., 2, 1896, 267 {Bacillus pseu- dotelanicus Migula, Syst. der Bakt., 2, 1900, 626; not Bacillus pseudotelanicus Kruse, idem; not Bacillus pseudoletanicus Chester, Man. Determ. Bact., 1901, 302; Bacillus pseudotelanicus var. aerobius Chester, ibid., 303); Plectridium palu- dosum Fischer, Jahr. f. Wiss. Bot., 27, 1895, 147; Pseudotetanicusbacillus, Tavel, Cent. f. Bakt., I Abt., 23, 1898, 538 {Bacillus pseudotetani Migula, Syst. der Bakt., 2, 1900, 598; Bacillus taveli Chester, loc. cit., 304). Also apparently identical with Bacillus sphaericus: Bacillus subietanicus Migula, Syst. der Bakt., 2, 1900, 629; Bacillus lactimorbus Jordan and Harris, Jour. Amer. Med. Assoc, 50, 1908, 1669 (see also Jour. Inf. Dis.,e, 1909,465) ; Bacillus serositidis Lacorte, Memorias do Instit. Oswaldo Cruz, 26, 1932, 1. There has been considerable confusion over the correct name to be applied to the non-pathogenic aerobic organisms re- sembling Clostridium tetani. Kruse {loc. cit.) isolated his cuture of Bacillus pseudoletanicus aerobius from a case of human tetanus. It was aerobic at ordi- nary temperatures but produced spores 728 MANUAL OF DETERMINATIVE BACTERIOLOGY only at higher temperatures and under anaerobic conditions. Migula called this Bacillus pseudotetanicus. Ford (Jour. Bact., 1, 1916, 520) stated that this name had priority over Neide's Bacillus sphaericus which he thought was iden- tical. On the other hand, Tavel {loc. cit.) isolated a pseudotetanus bacillus that was apparently anaerobic. Its spores were ellipsoidal and it formed more gas than the tetanus bacillus. Migula named this organism Bacillus pseudoteiani. Subsequentlj- both of Migula's names have been applied to the aerobic organism. Bacillus pseudo- tetanicus and Bacillus pseudotetani are nomina dubia and Bacillus sphaericus should therefore be used. Spores : Spherical, 0.7 to 1.3 microns in diameter, terminal to subterminal. Young spores in sporangia may be oval. Spore wall thick ; remnants of sporangium may adhere. Sporangia: Definitely swollen, clavate to spindle-shaped. Rods: 0.6 to 1.0 by 1.5 to 7.0 microns, occurring singly or in short chains. On glucose agar cells contain few small fat globules. Motile. Gram-variable; often Gram-negative with Gram-positive granules. Gelatin stab : Scant growth. No lique- faction. Gelatin hydrolyzed if tested by the Frazier technic. Agar colonies : Small, thin, flat, translu- cent, often spreading over the plate. Giant agar colonies : If the surface of the agar is fairly dry, many strains exhibit minute colonies that swarm out from the point of inoculation and cover the plate (of. Bacillus alvei and Bacillus circulans) . Agar slants: Growth thin, smooth, spreading, translucent, becoming yellow- ish-brown. Growth occurs at pH 6.0. Broth : Uniform turbidity. Glucose broth cultures have pH of 8.3 to 8.6 after 7 days. Milk : No change. Milk agar plate: Scant, if any, hydro- lysis of casein. Potato : Growth scant, thin, spreading, soft, gray, becoming yellowish-brown with age. Nitrites not formed from nitrates. Starch not hydrolyzed. No acid from carbohydrates. Acetylmethylcarbinol not produced. Citrates not utilized. Urease not formed. Salt tolerant. Growth occurs in broth containing 4 per cent NaCl. Optimum temperature about 30°C. Maximum temperature allowing growth 40°Cto45°C. Not pathogenic for guinea pigs. Aerobic, facultative. Source : From mud of a pond, rotting cypress wood, rotting oak wood, and from soil. Habitat : Widely distributed in nature. Bacillus rotans Roberts (Jour. Bact., 29, 1935, 229) differs from Bacillus sphaericus in that it will not grow as low as pH 6.0 nor in broth containing 4 per cent NaCl. Originally characterized by motile colonies, this phenomenon has been noted with certain other members of the genus and with some strains of Bacillus sphaericus. Smith, Gordon, and Clark {loc. cit.) consider it a variety of Bacillus sphaericus. Source : From intestine of a termite. Habitat: Probably widespread in soil. 18a. Bacillus sphaericus var. fusi- formis comb. nov. (Bacillus fusiformis Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 724.) From Latin/»SH.s, spindle ;/o?-m2s, shape. This organism differs from Bacillus sphaericus only in that it produces urease. Source : One strain isolated from Beta vulgaris lutea (beet). Also from milk, dust, soil and contaminated hirudin. Habitat : Widely distributed in nature. Bacillus loehnisii Gibson (Jour. Bact., 29, 1935, 495) is very closely related to the above. It will not grow at pH 6.0 or below, prefers media containing urea, and produces nitrites from nitrates. Gibson (loc. cit., 500) in discussing the FAMILY BACILLACEAE 729 organisms of this group stated "each species contains strains dissimilar in several features and each is connected to theothers by transitional forms". Smith, Gordon, and Clark {loc. cit.) tentatively have placed it as a variety of Bacillus sphaericus. Source : From soil. Habitat : Widely distributed in soil. 19. Bacillus pasteurii (Miquel) Ches- ter. ( Urobacillus pasteuTvi Miquel, Ann. Micrographie, 1, 1889, 552; sive Bacillus ureaea,ibid., 2, 1890, 13, 53, 122, 145, 367, 488; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 10, 1898, 110.) Named for the French scientist, Louis Pasteur (1822-1895). Viehover, Cent. f. Bakt., II Abt., 39, 1913, 209, gave the following as possible synonyms : Urobacillus maddoxii Miquel, Ann. Micrographie, 3, 1891, 27-5 and 305 {Bacterium maddoxi Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 98; Bacillus maddoxi Chester, ibid., 10, 1898, 110); Urobacillus leubei Beijerinck, Cent. f. Bakt., II Abt., 7, 1901, 51. Synonyms according to Gibson, Jour. Bact., 29, 1935, 494 and 496: Bacillus probatusYiehover, Cent. f. Bakt., II Abt., 39, 1913, 209; Urobacillus psychrocar- tericus and Urobacillus hesmogenes Ru- bentschik. Cent. f. Bakt., II Abt., 66, 1925, 166 (Bacillus psychrocartericus and Bacillus hesmogenes Bergey et al., Man- ual, 3rd ed., 1930, 403, 404). Gibson also included the following as possibly identi- cal with the above although they were incompletely described: Bacillusureae II and /// Burri, Herfeldt and Stutzer, Jour. Landw., 4^, 1894, 329; Urobacillus duclauxii Miquel, Ann. Micrographie, 2, 1890, 53, 122, and 145; Urobacillus mad- doxii Miquel, ibid., 3, 1891, 275 and 305. Description taken from Lohnis and Kuntze, Cent. f. Bakt., II Abt., 20, 1908, 684 ; Gibson, Jour. Bact., 28, 1934, 295 and 313 ; Smith, Gordon, and Clark {loc. cit.). This species has been designated as the type species of the genus Urobacillus Miquel (Ann. de. Micrographie, 1, 1889, 517) by Enlows (U. S. Pub. Health Ser., Hyg. Lab. Bull. 121, 1920, 96). Spores: Spherical, 1.0 to 1.2 microns, terminal to subterminal. Sporangia: Prevailingly clavate. Not in chains. Rods: 0.7 to 0.8 by 1.5 to 2.0 microns (1.0 to 1.5 by 4.0 to 5.0 microns, Lohnis and Kuntze), occurring singly or in pairs. Motile. Gram-variable. Urea gelatin stab : Slow liquefaction. Urea agar colonies: Small, entire, not characteristic. Urea agar slope: Growth thin, very little spreading, colorless or white to yellowish. Will not grow at pH 6.0 or less. L^rea broth: Moderate to heavy uni- form turbidity. Will grow with 4 per cent NaCl added. Nitrites produced from nitrates in urea nitrate nutrient broth. Starch not hydrolyzed. Carbohydrates not attacked. Acetylmethylcarbinol not formed. LTrease produced. Optimum temperature about 30°C, minimum 6°C. Maximum temperature allowing growth 40°C in water bath. Optimum temperature for urease activity 50°C. Aerobic. The distinguishing character of this species is that growth occurs only in peptone media containing urea or free ammonia under neutral or alkaline conditions. Source: From decomposing urine. Habitat: Widely distributed in soil, dust, manure, and sewage. 20. Bacillus thermoamylolyticus Cool- haas. (Cent. f. Bakt., II Abt., 75, 1928, 344.) From Greek thermos, hot, amylon, starch, and lytikos, able to loose; hence, dissolving. Probably intended to mean thermophilic and starch digesting. Spores: Slightly elongated, ellipsoids 0.6 by 1.5 microns, central. Sporangia: Cylindrical, not swollen, not in chains. 730 MANUAL OF DETERMINATIVE BACTERIOLOGY' Rods: 0.6 by 5 to 8 microns. Motile. Gram-positive. Gelatin stab : No liquefaction. Agar colonies: At 60°C of two types, large and small, circular, translucent, granular, slimy. Broth: Very weak growth, no surface growth, no sediment. Milk: Not coagulated, slowly pepton- ized. Potato : Slight growth. Nitrites produced from nitrates. Starch actively hydrolyzed. Acid and gas from glucose, fructose, galactose, maltose, dextrin, starch and glycerol. Arabinose, xylose and manni- tol not fermented Thermophilic, optimum temperature 50° to 55°C. Aerobic. Source: From sewage. Habitat : Probably in decaying matter. 21. Bacillus kaustophilus Prickett. (N. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928, 38.) From Greek kaustos, burnt, red-hot; philos, loving; heat-loving. Spores: Ellipsoidal, 0..'3 by 0.6 to 0.8 micron, terminal to subterminal. Xo free spores observed. Sporangia: Only slightly swollen if at all. Rods: On yeast extract -nutrient agar at 56°C, 0.7 by 2.0 to 4.5 microns, with rounded ends. Actively motile. Gram- positive. Gelatin stab: No growth at 20°C. Liquefied at 56°C. Agar colonies: At 56°C, circular, con- vex, smooth. Borders entire to irregular. Show curled structure, strands of chains. Brown by transmitted light. Agar slant : Growth abundant, raised, glistening, contoured, bluish-green to bluish-white by transmitted light. After three weeks at 37°C, growth has a distinct reddish-brown color, butyrous, viscid. Broth: Slightly turbid, no sediment. No surface growth, alkaline. Litmus milk: Rennet coagulum, pep- tonization feeble, litmus reduced. Potato : Amount of growth Variable, brownish, spreading, glistening, slimy. Some strains do not grow. Nitrites produced from nitrates, often with the production of nitrogen. Starch is hydrolyzed. Acid from glucose and salicin. Rham- nose, maltose, sucrose, raffinose, manni- tol, sorbitol and inulin not fermented. Acetylmethylcarbinol not produced. Thermophilic, optimum temperature 60°C to 65°C. Growth at 73°C to 75°C but none at 80°C on agar slants. Aerobic, facultative. Good growth occurs in synthetic media containing potassium nitrate, sodium ammonium phosphate, aspartic acid, and sodium asparaginate, respectively, as only sources of nitrogen with glucose as source of carbon. Source : Forty -eight cultures isolated from pastQurized milk at a single milk plant (Buffalo, N.Y.). Habitat : Probably originally from soil and dust. Thermobacillus digestans Feirer (Soil Sci., 23, 1927, 50) seems to be closely related to the preceding. It is more strongly proteolytic, digesting milk com- pletely in 7 days. Source : Four strains isolated from soil. 21a. Bacillus pepo Shaw. (Jour. Inf. Dis., 43, 1928, 473.) From Cucurbita pepo, the pumpkin. From the brief original description, this organism seems to vary from Bacillus kaustophihts only in its distinctive viscid or slimy character. Source: Two cultures isolated from swelled cans of pumpkin. Habitat : Probably found in soil and dust. 22. Bacillus thermoindiflferens Wein- zirl. (Jour. Med. Research, 39, 1919, 402.) From Greek thermos, hot and Latin indifferens, indifferent. Indiffer- ent to or tolerant of heat. Spores: Ellipsoidal, 0.5 by 0.8 micron, terminal. FAMILY BACILLACEAE 731 Sporangia: Cylindrical, not swollen. Rods : 0.7 by 2.0 to 4.5 microns, occur- ring singh- and in short chains, with rounded ends. ^lotile. Gram-positive. Gelatin stab: Growth filiform. Slow infundibuliform liquefaction. Agar colonies : Circular, convex, smooth, entire, amorphous. Agar slant: Growth flat, spreading, glistening, translucent, butyrous, con- toured . Broth: Turbid, abundant sediment. Xo surface growth. Litmus milk: Alkaline. Litmus reduced. Potato : No growth. Indole not formed. Nitrites not produced from nitrates. Starch is hydrolyzed. Acid froni glucose. No acid from lactose, sucrose or mannitol. Thermophilic, optimum temperature 55°C. Grows at 20° to 37°C. Aerobic . Source : Isolated from canned pumpkin. Habitat : Probably found in soil and dust. Thermobacillus reductans Feirer (Soil Sci., 23, 1927, 51) is said to resemble Bacillus thermoindifferens except that nitrites are formed from nitrates and the minimum temperature is 40°C. Source: Two strains isolated from soil. Thermobacillus catenatus Feirer (Soil Sci., 23, 1927, 53) may be related to this group. The description is very incom- plete. Its distinctive feature is the production of indole. Source : Two strains isolated from soil. 23. Bacillus thermodiastaticus Ber- ge}- et al. (Type 1, Jiergey, Jour. Bact., 4, 1919, 304; Bergey et al.. Manual, 1st ed., 1923, 310.) From Greek thermos, hot and diaslalikos, separative; M. L., enzjaiiatic, diastatic; hence diastatic at high temperatures. Spores : Of less diameter than that of the rods, ellipsoidal, central. Sporangia: Cj'lindrical. Rods: 0.5 to 0.7 by 2 to 3 microns, occurring in chains, with square ends. Motile with peritrichous flagella. Gram- positive. Gelatin stab : Liquefaction. Agar colonies : Grayish, spreading, with lobate to funbriate borders. Agar slant: Growth thin, limited, bluish-gray. Broth: Turbid. Litmus milk: Not coagulated, pep- tonized. Potato: Growth slight, grayish. Nitrites produced from nitrates. Starch is hydrolyzed. Thermophilic, optimum temperature 65°C. No growth at 50°C. Growth at 75°C. Aerobic . Source: Isolated from dust and con- taminated milk. Habitat : Probably widely distributed in soil and dust. Thermobacillus diastasius Feirer (Soil Sci., 23, 1927, 49) differs from Bacillus thermodiastaticus only in that nitrites are not formed from nitrates (Feirer). Source : Two strains isolated from soil. 24. Bacillus cylindricus Blau. (Cent, f. Bakt., II Abt., 15, 1905, 119.) From Greek kylindrikos, cylindrical. Spores : Somewhat elongated, 0.7 to 1.1 by 1 .8 to 2.5 microns, terminal. Remnants of sporangium adherent. Germination equatorial and oblique. Sporangia : Cylindrical or only slightly swollen at end, not in chains. Rods : On glucose agar at 60°C, 0.8 to 1.1 by 5.0 to 7.5 microns, occurring singly and in pairs. Motile with peri- trichous flagella. Cells store glycogen. Gram-positive. Gelatin stab : No liquefaction. Glucose agar colonies : Grayish-white, entire to lobed to dentate. By trans- mitted light yellowish-brown centers with brownish-yellow borders. Finely fibrous structure. Glucose agar slant : Growth thin, dull, grayish-white. Litmus milk : Unchanged. 732 MANUAL OF DETERMINATIVE BACTERIOLOGY Potato : No growth. Nitrites not produced from nitrates. Starch not hydrolyzed. Thermophilic, optimum temperature 60°C to 70°C. Source : Isohxted from moist field soil in Germany . Habitat : Probably found in dust and soil. Bacillus calidus Blau (Cent. f. Bakt., II Abt., 15, 1905, 134) differs so little from the preceding species that it cannot be considered as distinct. Source : Isolated from field soil in Germany (Blau). Dust, ground feeds, etc., about dairies and various dairy products (Prickett, N. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928,45). 25. Bacillus robustus Blau. (Cent. f. Bakt., II Abt., 15, 1905, 126.) From Latin robustus, oaken, hard, firm. Spores: Ellipsoidal, 1.0 by 1.6 to 2.2 microns, polar to medial. Remnants of sporangium not adherent. Germination prevailingly equatorial. Sporangia: Ellipsoidal to cylindrical, not in chains. Rods: 1.0 to 1.2 by 3 to 4 microns, occurring singly and in short chains. Motile. Gram-positive. Glucose agar colonies : Circular, gray- white. By transmitted light brownish- yellow. Borders distinct, serrate to lobed, finely granular. Glucose agar slant : Growth yellowish- white, translucent, becoming grayish- white, spreading, dull. Potato : Growth yellowish-whitc, moist, glistening, smooth. Nitrites not produced from nitrates. Starch not hydrolyzed. Thermophilic, optimum temperature 55°C to 60°C. Grows at 65°C. Aerobic . Source : Isolated from field soil near a forest in Germany. Habitat : Probably found in soil and dust. Thermobacillus restatus Feirer (Soil Sci., 23, 1927, 51) is said to correspond in some respects with Bacillus robustus. Feirer states that it is not possible to definitely establish the identity because Blau failed to record the action of Ba- cillus robustus on nitrates and several other media and did not note the produc- tion of H,S. Source : Three strains isolated from soil. 25a. Bacillus losanitchii Bergey et al. {Bacillus thermophilus losanitchii Georgevitch, Cent. f. Bakt., II Abt., 27, 1910, 164; Bergey et al., Manual, 1st ed., 1923, 313.) Named for Losanitch, near Vranje. As far as can be determined from the meager description, this organism does not differ from Bacillus robustus except perhaps as to the maximum temperature allowing growth. Growth limits are 45°C to 78°C. Source: Isolated from water of hot sulfur spring. Temperature of water 83°C. Habitat : Probably in natural hot waters. Note: Georgevitch (Arch. f. Hyg., 72, 1910, 201) has described a thermophilic aerobic spore-forming sulfur bacillus from a hot sulfur spring at Vranje (Ser- via) under the name Bacillus thermo- philus vranjensis. This does not grow on ordinary media unless sulfur com- pounds are added. It has a tuft of flagella at either end. Spores are ellipsoidal, terminal, distend the rod, and show polar germination. Georgevitch (Cent. f. Bakt., II Abt., 27, 1910, 150) describes a second thermo- philic, motile, capsulated, ellipsoidal- spored rod from a chalybeate hot spring near Vranje under the name Bacillus t h ermoph il us jivo in i . 26. Bacillus calidolactis Hussong and Hammer. (Jour. Baet., 15, 1928, 186.) From Latin calidus, warm, hot and lac, lactis, milk. Gorini states (R. 1st. Lombardo Sci. e Lett. Rend., 76, 1942, 3) that Bacillus FAMILY BACILLACEAE 733 calidolaciis is the same organism as Bacillus lactis termophilus (sic) Gorini (Giorn. d. R. Soe. Ital. d'Igiene, 16, 1894, 16). From the descriptions this appears to be probable. Spores: Ellipsoidal, of slightly greater diameter than the rods, terminal. Sporangia: Slightly swollen, clavate. Rods : 0.7 to 1.4 by 2.6 to 5.0 microns, occurring singly, in pairs and short chains. Non-motile. Gram-positive, some cells becoming Gram-negative with age. Gelatin stab: Xo liquefaction. No growth on plain nutrient agar. Glucose agar colonies: Thin, white, opaque, filamentous. Glucose agar slant : Growth abundant, echinate, dull, white. Glucose agar stab: Growth abundant, beaded, gray. Glucose broth: Turbid. Litmus milk: Acid, coagulation. Lit- mus reduced. Potato: No growth. Nitrites produced from nitrates by some strains. Acid from glucose, lactose, fructose, galactose and maltose. No acid from inulin, sucrose or glycerol. Thermophilic, optimum temperature 55=C to 65°C. No growth at 37°C. Aerobic, facultative. Source: Isolated from normal pasteur- ized skim milk (Hussong and Hammer). Milk and milk powder (Prickett, N. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928, 47). Habitat : Probably in dairy products. 27. Bacillus michaelisii Prickett. {Bacillus thennophilus aquatilis lique- faciens Michaelis, Arch. f. Hyg., 36, 1899, 285; Prickett, N. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928, 45.) Named for Georg Michaelis of Berlin who first isolated the species. Spores : Of greater diameter than the rods, terminal. Sporangia: Swollen, clavate. Rods: 0.6 to 0.8 by 2 to 4 microns. Motile. Gram -positive. Gelatin stab : Liquefaction. Agar colonies : Circular, raised, smooth, glistening. Agar slant : Growth moderate, smooth, glistening. Broth: Slight turbidity. Milk: Not coagulated, alkaline. Potato : Growth moist, glistening, yel- lowish, becoming brownish. Nitrites with gas produced from nitrates. Starch is hydrolyzed. Acid from glucose and sucrose. No acid from rhamnose, maltose, lactose, glycerol, mannitol or inulin. Thermophilic, optimum temperature 50°C to 60°C. Aerobic, facultative. Source : Isolated from fountain waters (Michaelis). From fodder, dust, dairy utensils (Prickett). Habitat : Probably found in soil and dust. 27a. Bacillus lohatus Bergey et al. (Type 3, Bergey, Jour. Bact., I^, 1919, 304 ; Bergey et al., Manual, 1st ed., 1923, 311.) From Greek lohatos, having the form of a lobe. Judging from the meager description, there is no essential difference between this organism and the preceding. Source: Isolated from dust, soil, and horse manure. Habitat : Probably widely distributed in soil and decaying matter. Bacillus nondiastalicus Bergey et al. (Type 2, Bergey, Jour. Bact., 4, 1919, 304 ; Bergey et al.. Manual, 1st ed., 1923, 310.) From Greek, no diastase. The description of this organism is practically identical with Bacillus loba- tus, the only difference noted being that this species hydrolyzes starch while Bacillus nondiastalicus does not. Source : Isolated from dust and soil (Bergey). Ground grains, raw and pas- teurized milk (Prickett, N. Y. Agr. E.xp. Sta. Tech. Bui. 147, 1928, 47). Thermobacillus vulgaris Feirer (Soil Sci., ^3, 1927, 50) liquefies gelatin, does 734 MANUAL OF DETERMINATIVE BACTERIOLOGY not reduce nitrates to nitrites nor alter litmus milk. According to Feirer it is otherwise similar to Bacillus nondiasta- ticus. Source : Two strains isolated from soil . 27b. Bacillus tlierinunouliquefaciens Bergey et al. (Type 4, Bergey, Jour. Bact., 4, 1919, 304; Bergey et al., Manual, 1st ed., 1923, 312.) From Greek thermos, hot; and Latin non, not and liquefaciens, making liquid. Probably intended to mean thermophilic and non-gelatin- liquefying. Aside from the non-liquefaction of gelatin, there seems to be no difference in the description of this organism and the two immediately preceding. Source: Isolated from dust, soil, and horse manure. Habitat: Probably found in soil and decaying matter. 28. Bacillus thermotranslucens Bergey et al. (Type 5, var. b, Bergey, Jour. Bact., 4, 1919, 304; Bergey et al.. Manual, 1st ed., 1923, 312.) From Greek thervws, hot and Latin irausluceits, translucent. Probably intended to mean thermophilic and translucent. Spores: Of larger diameter than the rods, terminal. Sporangia : Terminally swollen, clavate, not in chains. Rods : 0.3 to 0.4 by 1.0 to 1.5 microns, occurring singly. Motile with peritrich- ous flagella. Gram-positive. Gelatin stab: No liquefaction. Agar colonies: Thin, transparent, spreading widely. Agar slant: Growth thin, spreading, veil-like. Broth: Turbid. Litmus milk: Not coagulated, slightly acid. Potato: No growth. Nitrites not produced from nitrates. Starch slightly hydrolyzed. Thermophilic, optimum temperature 60°C. Slight growth at 37°C. No growth at 70°C. Aerobic . Source: Isolated from guinea pig feces, dust and from cheese. Habitat : Probably found in soil and decaying matter. Thermohacillus linearius Feirer (Soil Sci., 23, 1927, 53) is said to be similar in some respects to the preceding. Feirer states that formation of acid from several sugars is the distinctive feature of this species, a character not mentioned by Bergey. Source: Five strains isolated from soil. 2Sa. Bacillus slearothermophilus Donk. (Jour. Bact., 5, 1920, 373.) From Greek sledr, tallow and thermofhilos, heat- loving. Intended meaning obscure. From the descriptions, the vegetative rods of this organism seem to be slightly larger than the preceding, otherwise no difference is noted. Source : Isolated from samples of spoiled canned corn and string beans. Habitat: Probably found in soil and dust. 28b. Bacillus aerothermophilus Wein- zirl. (Jour. Med. Research, 39, 1919, 403.) From Greek aeros, air and /';e?'- mophihis, heat-loving. Probably in- tended to mean aerobic and thermophilic. There is nothing in the original account of this organism which is at variance with that of the preceding. Source : Isolated from canned string beans (Weinzirl). From milk, water, hay, dust, beef extract, and agar (Prick- ett, N. Y. Agr. Exp. Station Tech. Bull. 147, 1928,46). Habitat : Probably found in soil and dust . Thermobacilhis alcalinus Feirer (Soil Sci., 23, 1927, 52) is said to differ from the preceding in that it does not change litmus milk. Source : Four strains isolated from soil. Thermobacillus ruber Feirer (Soil Sci., 23, 1927, 52) apparently is closelj' related to this group. Its distinguishing char- acter is the production of a pink pigment FAMILY BACILLACEAE 735 in meat, brain, and blood serum, no color on other media. Source : Isolated from soil. 29. Bacillus thermocellulolyticus Cool- haas. (Cent. f. Bakt., II Abt., 76, 1928, 43.) From Greek thermos, hot; and Latin cellula, a small room; M. L., cellu- lose and Greek lytikos, dissolving. Prob- ably intended to mean thermophilic and cellulose-digesting. Spores: Ellipsoidal, 0.8 by 1.5 microns, terminal. Sporangia : Terminally swollen, clavate. Rods : 0.3 by 3.5 to 6 microns, occurring singly and in pairs . Xo reserve material . Non-motile. Gram-positive. Gelatin stab: Xo liquefaction. Glucose agar colonies : Small, glisten- ing, translucent. Cellulose agar colonies : Circular, bor- ders undulate to lobate. Broth : Slight growth, no surface growth or sediment. Milk: No growth. Nitrites not produced from nitrates. Starch is hydrolyzed. No acid from carbohydrates. Cellulose hydrolyzed. Thermophilic, optimum temperature 50°C to 55°C. Maximum 60°C to 65°C. ^Minimum 35°C to 37°C. Aerobic, facultative. Source: Isolated from sewage. Habitat : Probably found in decaying matter. 30. Bacillus thermoalimentophilus Weinzirl. (Jour. Med. Research, 39, 1919, 404.) From Greek thermos, hot; Latin alimentum, food; and Greek ■philos, loving. Loving hot food. Spores: Ellipsoidal, 0.8 by 1.0 micron, terminal. Sporangia: Swollen, clavate, not in chains. Rods: 0.6 by 3.0 microns, occurring singly, with rounded ends. Motile, flagella not stated. Gram-positive. Gelatin stab : No growth at 20°C. Agar colonies : Circular, raised, smooth, amorphous, entire. Agar slant : Growth spreading to effuse, smooth, glistening, butyrous. Broth: Turbid, surface ring. Litmus milk: Unchanged. Potato : Xo growth. Nitrites with gas produced from ni- trates. Starch not hydrolyzed. Neither acid nor gas from glucose, lac- tose, sucrose or raannitol. Thermophilic, optimum temperature 55°C. Xo growth at 20°C. Growth at 37°C. Aerobic. Source : Isolated from canned blueber- ries (Weinzirl). Pasteurized milk and filter cloth (Prickett, X. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928, 46). Habitat : Probably found in soil and dust. ThermubaciUus violaceus Feirer (Soil Sci., 33, 1927, 52) corresponds in some respects with the preceding. Feirer also states that his cultures did not reduce nitrates to nitrites and produced acid on glucose and sucrose. Source : Four strains isolated from soil. 31. Bacillus thermoliquefaciens Ber- gey et al. (Type 5, var. a, Bergey, Jour. Bact., 4, 1919, 304 ; Bergey et al., Manual, 1st ed., 1923, 313.) From Greek thermos, hot, and Latin liquefaciens, liquefying. Probably intended to mean thermophilic and gelatin-liquefying. Spores: Ellipsoidal, polar, of greater diameter than the rods. Sporangia : Terminally swollen, clavate. Rods: 0.2 to 0.4 by 2 to 3 microns, occurring singly, with rounded ends. Motile with peritrichous flagella. Gram- positive. Gelatin stab : Liquefaction. Agar colonies : ^Moderately dense, lobate. Agar slant : Growth dense, grayish, lobate to fimbriate margins. Litmus milk: Coagulated, acid. Lit- mus reduced. 736 MANUAL OF DETERMINATIVE BACTERIOLOGY Potato : No growth. Nitrites and ammonia produced from nitrates. Starch not hydrolyzed. Thermophilic, optimum temperature 60°C. Slight growth at 37°C. No growth at 70°C. Aerobic . Source: Isolated from dust, guinea pig feces and horse manure (Bergey ) . Water and milk (Prickett, N. Y. Agr. Exp. Sta. Tech. Bui. 147, 1928, 47). Habitat : Probably originally from soil ■ nd dust. 32. Bacillus tostus Blau. (Cent. f. Bakt., II Abt., 15, 1905, 130.) From Latin tostus, parched, dried. Spores: Ellipsoidal, 0.8 to 1.6 by 1.5 to 2.2 microns. Germination prevail- ingly polar. Sporangia : Terminally swollen, clavate, not in chains. Rods : 1.2 by 4.5 to 5.0 microns, occur- ring in pairs and in short chains. Cells store glycogen. Motile with peritrichous flagella. Agar colonies: Small, circular, dense. By transmitted light bright yellow to yellowish-brown. Borders sharp, entire to lobate. Older colonies porcelain-like. Agar slant : Growth thin, grayish-white, spreading, smooth, glistening. Potato : No growth. Nitrites not produced from nitrates. Starch is hydrolyzed. Ammonia is produced. Thermophilic, optimum temperature 60°C to 70°C. Aerobic. Source: Two cultures isolated from soil in Germany. Habitat: Probably found in soil and dust. 33. Bacillus viridulus (Migula) Bergey et al. {Bacillus thermophihis II Rabino- witsch, Ztschr. f. Hyg., 20, 1895, 154; Bacterium viridulum Migula, Syst. der Bakt.,;g, 1900, 343; Bacterium thermophi- lum 7/ Chester, Manual, 1901,186; BaciZ- lus thermophilus Bergey et al.. Manual, 1st ed., 1923, 315; Bergey et al.. Manual, 4th ed., 1934, 464; not Bacillus thermo- philus MiqueL Ann. d. Microg., 1, 1888, 6; not Bacillus thermophilus Chester, Man. Determ. Bact., 1901, 265.) From Latin, dim. adj. viridis, green, some- what green. Spores: Si)herical, central. Sporangia: Cylindrical, not swollen. Rods : Rather large, slightly bent, occurring singly and in pairs. Non- motile. Gram-positive. Agar colonies : Irregular, spreading, granular, greenish. Broth: Alkaline. Potato : Growth grayish-yellow ; margin undulate. Nitrites not produced from nitrates. Starch is hydrolyzed. Thermophilic, grows at 60°C to 70°C. Optimum temperature 62°C. Grows at 33°C. Aerobic, facultative. Source : Isolated from soil, snow, feces, corn grains. Habitat : Probably found in soil and dust. Appendix : The following additional aerobic spore-forming bacteria are found in the literature. Because of insufficient data it has not been possible to classify them. Some of these may be synonyms of well-known species, some may be varieties, whereas others may actually be separate species. Aromabacillus weigmanni Omeliansky. (Isolated by Weigmann, 1890; Omelian- sky, Jour. Bact., 8, 1923, 398.) From milk. Bacillus abysseus ZoBell and Upham. (Bull. Scripps Inst, of Oceanography, Univ. Calif., 5, 1944, 273.) Subterminal spores. From marine mud. Bacillus acidifaciens Patrick and Werkman. (Iowa State Coll. Jour. Sci., 7, 1933, 413.) One of a group charac- terized by the fermentation of xylan. A single culture isolated from decayed maple wood. Bacillus acidificans presamigenes casei FAMILY BACILLACEAE 737 Gorini. (Rend. R. Accad. dei Lincei, 8, 1928, 598.) From manure, fodder and milk. Regarded by Gorini (personal communication, 1925) as identical with Bacillus circulans Jordan. Bacillus acido-proteolyticus casei Gorini. (Le Lait, 9, 1912, 98.) From Parmesan and Emmenthal cheese. Re- garded by Gorini (personal communica- tion, 1925) as equivalent to one of the species of Tyrothrix of Duclaux. Bacillus adametzi Trevisan. (Brauner Wasserbacillus, Adametz and Wichmann, Mittheil. d. oesterr. Versuchsstat. f. Brauerei u. Malzerei, Wien, Heft 1, 1888, 51 ; Trevisan, I generi e le specie delle Batteriacee, 1889, 19; not Bacillus adametzii Migula, Syst. d. Bakt., 2, 1900, 686; Bacillus brunneus Eisenberg, Bakt. Diag., 3 Aufl., 1891, 142; Bacterium bruii- ueurn JMigula, ibid., .331; not Bacterium brunneneum Schroeter, in Cohn, Beitr. z. Biol. d. Pflanz., ;, Heft 2, 1882, 125.) From water. Bacillus adhaerens Laubach. (Jour. Bact., 1, 1916, 503.) One culture iso- lated from dust. Bacillus aegypiiacusWerner . (Cent. f. Bakt., II Abt., 87, 1933, 459.) Good growth on Ca n-butyrate agar. One culture isolated from Egyptian soil. Bacillus aerifaciens Steinhaus. (Jour. Bact., 42, 1941, 782.) Author states that it probably belongs to the Aerobacillus group. From triturated specimens of the white cabbage butterfly (Pieris rapae). Bacillus aerobius von Wahl. (Cent. f. Bakt . , II Abt .,16, 1906, 496. ) Reported to resemble Bacillus mesentericus fuscus . From canned peas. Bacillus aerophilus Fliigge. (Fliigge, Die Mikroorganismen,2 Aufl., 1886, 321; Bacterium aerophilum Chester, Man. De- term. Bact., 1901, 191.) From dust. Bacillus afanassieffi Trevisan. {Ba- cillus tussis convtdsivae Afanassief, St. Petersburg, med. Wochnschr., 1887, No. 38-42; not Bacillus tussis convulsivae Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 269; Trevisan, I generi e le specie delle Batteriacee, 1889, 13; Ba- cillus pertussis Migula, Syst. d. Bakt., 2, 1900, 754.) From mucus and pus. Bacillus agilis Tschistowitsch. (Tschistowitsch, Berl. klin. Wochnschr., 1892, 512; not Bacillus agilis Trevisan, I generi e le specie delle Batteriacee, 1889, 14; not Bacillus agilis Chester, Man. Determ. Bact., 1901, 226; not Bacillus agilis Mattes, Sitzungsber. d. Gesellsch. z. Beforderung d. gesam. Naturw. z. Marburg, 62, 1927, 406; not Bacillus agilis Hauduroy et al.. Diet. d. Bact. Path., Paris, 1937, 33.) From pus. Bacillus agilis Hauduroy et al. {Ba- cillus agilis larvae Toumanoff, Bull. Soc. Cent, de Med. V^ter., 80, 1927, 367; Hauduroy, Ehringer, Urbain, Guillot and Magrou, Dictionnaire de Bact^ries Pathogenes, Paris, 1937, 33.) Found in foulbrood of bees. Bacillus agrestis Werner. (Cent. f. Bakt., II Abt., 87, 1933, 468 ; not Bacillus agrestis de Rossi, Microbiol, agraria e technica, Torino, 1927, 828.) One of a group of species described as being able to grow on a Ca n-butyrate agar. Three cultures were isolated from German and Italian soils. Bacillus agri Laubach and Rice. (Jour. Bact., 1, 1916, 516.) Isolated twice from soil in Baltimore. Bacillus agrophilus Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 189.) Only moderate growth on Ca n-butyrate agar. One culture isolated from soil from Cuba. Bacillus agrotidis typhoides Pospelov. (Rept. Bur. Appl. Ent., Russian, 3, 1927, 8.) Found in diseased larvae of the moth, Euxoa (Agrotis) segctum. Bacillus {Streptobacter) albuminis Schroeter. (Bacillus aus Faeces V, Bien- stock, Ztschr. f. klin. Med., 8, Heft 1, 1884, 1 ; Schroeter, in Cohn, Kryptog. Flora V. Schlesien, 3, 1, 1886, 162; Ba- cillus putrificus coli Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 303; Bacillus diaphthirus Trevisan, I generi e le specie delle Batteriacee, 1889, 15.) From feces. Bacillus albus (Sack) Bergey et al. {C elhdomonas albus Sack, Cent. f. Bakt., II Abt., 62, 1924, 79; Bergey et al., Man- 738 MANUAL OF DETERMINATIVE BACTERIOLOGY ual, 3rd ed., 1930, 398; not Bacillus ulbus Trevisan, I generi e le specie delle Batteriacee, 1889, 14; not Bacillus albus Copeland, Report Filtration Comm., Pittsburgh, 1899, 344.) Cellulose is hydrolyzed. From soil in GermanJ^ Bacillus alcalophilus Vedder. (Ned. Tijdschr. v. Hyg. Microbiol, en Serolog., 1, 1934, 141.) Grows only in and on highly alkaline culture media. Sixteen strains isolated from the feces of healthy animals. Bacillus alopecuri Nogtev. (Botani- cheskii Zhurnal, U.S.S.R., 23, 1938, 149.) Causes nodule formation on fox grass {Alopecurus pratensis). Bacillus alpha Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 366.) From the air. Bacillus alpinus Werner. (Cent. f. Bakt., II Abt., 87, 1933, 465.) Good growth on calcium salts of formic, acetic, propionic and butyric acids. One cul- ture isolated from soil from Austria. Bacillus alveolaris Ksenjoposky. (Re- view of pests of Volhymia, Volhymia Ent. Bur., Zemstvo of Volhymia, Zitomir, 1916, 24 pp.) From diseased bees {Apis mellifera). Bacillus amarus Hanmier. (Iowa Agr. Exp. Station Res. Bull. 52, 1919, 198.) From evaporated milk. Bacillus aminovorans den Dooren de Jong. (Cent. f. Bakt., II Abt., 71, 1927, 215.) From soil. Bacillus amyloaerohius Crimi. (Abst. in Cent. f. Bakt., II Abt., 61, 1924, 63.) From potato rot. Bacillus amylolyticus Kellerman and McBeth. (Cent. f. Bakt., II Abt., 34, 1912, 490.) Decomposes cellulose. One culture isolated from manure. Bacillus annuliformis Migula. (Fa- denahnlicher Bacillus, Maschek, Bakt. Untersuch. d. Leitmeritzer Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt, 2, 1900, 783.) From water. Bacillus anthracis similis McFarland. (Cent. f. Bakt., I Abt., 24, 1893, 556.) From dust. Bacillus apicum Canestrini. (Atti Soc. Ven. Trent. Sci. Nat., 91 ; according to Kruse, in Fliigge, Die Mikroorganis- men, 3 Aufl., 2, 1896, 233.) From dis- eased bees and their larvae. Bacillus aporrhoeus Fuller and Nor- man. (Jour. Bact., 4^, 1943,277.) From soil. Decomposes cellulose. Bacillus arachnoideus Migula. (Bacil- lus No. Ill, Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lactis No. Ill, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 208; Migula, Syst. d. Bakt., 2, 1900, 583; Bacterium lacteum Migula, ibid., 321.) From milk. Bacillus arenarius Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 187.) Good growth on Ca n-butyrate agar. One strain isolated from Cuban soil. Bacillus aridus Migula. (Bacillus No. 8, Pansini, Arch. f. path. Anat., 122, 1S90, 444; Migula, Syst. d. Bakt., 2, 1900, 559.) From sputum. Bacillus arlongii (sic) DeToni and Trevisan. (Bacillus de la septicemie gangreneuse, Arloing and Chauveau, see Crookshank, Man. of Bact., 3rd ed., 1890, 305; DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 950.) From wounds in gangrenous septicaemia. Bacillus asiaticus Sakharoff. (Sak- haroff, Ann. Inst. Past., 8, 1893, 550; not Bacillus asiaticus Castellani, Cent. f. Bakt., I Abt., Orig., 65, 1912, 262.) From feces in a case of cholera. Bacillus asteris Verona. (Riv. Pat. Veg., 25, 1935, 15.) Pathogenic for China aster (Aster chinensis). Bacillus asthenogenes Bernard. (Ann. Inst. Past., 35, 1921, 459.) Grows aero- bically as well as anaerobically. Under anaerobic conditions it is said to play a role in gastric derangement and infection commonly confused with beriberi. Author reports that it is very similar to Bacillus megatherium. Bacillus aterrimus tschitensis Kli- menko. (Cent. f. Bakt., II Abt., 20, 1908, 1.) Reported to be like the black potato bacillus except that it forms a black pigment on gelatin and the potato is brown instead of black. From air. Bacillus aurantius (Sack) Bergey etal. FAMILY BACILLACEAE 739 (Cellulomonas aurantius Sack, Cent. f. Bakt., II Abt., 62, 1924, 78; Bergey et al., Manual, 3rd ed., 1930, 421; not Bacillus aurantius Trevisan, I generi e le specie delle Batteriacee, 1889, 19.) From soil. Bacilhis badius Batchelor. (Jour. Bact., 4, 1919, 25.) From the intestinal tract of children. Bacilhis balcamis Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 25.) Growth on media containing m/50 phenol. Eight strains isolated from soil. Bacillus barbitistes Statelov. (Mitt. bulg.ent.Gesells., 7, 1932,56-61.) From diseased tettigonids {Isophya (Barbi- tistes) aynplipennis) . Bacillus batatae Otani. (Trans. Tot- tori Soc. Agric. Sci., Japan, 6, 1939, 222.) From rotten sweet potatoes {Ipomaea batatas). Bacillus bellus Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 96.) Probably a strain of Bacillus brevis. One culture isolated from garden soil of Germany. Bacillus bernensis Lehmann and Neu- mann. (Aromabacillus, Burri, Cent. f. Bakt., II Abt., 3, 1897, 609; Lehmann and Neumann, Bakt. Diag., 2 Aufi., 2, 1899, 304 ; Bacillus odoratus Migula, Syst. d. Bakt., ^, 1900, 686; Bacterium odoratum Omeliansky, Jour. Bact., 8, 1923, 394.) From Emmenthal chee-se. Reported as producing the aroma of this cheese. Bacillus beta Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 366.) From dust. Bacillus betainovorans Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 94.) Good growth on betaine and valine agar. One culture isolated from soil from Mantua. Bacillus beianigrificans Cameron, Esty and Williams. (Food Research, 1, 1936, 75.) From blackened canned beets where juice contains an abnormally high amount of iron. Bacillus biacutum Soriano. (Revista del Inst. Bacteriol., Univ. Buenos Aires, 6, 1935, 564.) From soil. Bacillus bombycis Macchiati. (Des vibrions, Pasteur, Etudes sur la maladie des vers a soie, La Flacherie, Chapitre II, Paris, 1870; Macchiati, Stazioni sperimentali Agrarie Italiane, 20, 1891, 121; not Bacillus bo7nbycis Chatton, Compt. rend. Acad. Sci., Paris, 156, 1913, 1708; Bacillus viegaterium bombycis Saw- amura, Tokyo Imp. Coll. Agric. Bull., 6, 1905, 375.) Pasteur originally isolated this large bacillus from the intestine of silkworms {Bombyx mori) suffering from wilt disease. Regarded by Sawamura as a variety of Bacillus megatherium. Bacillus bombycis non-liquefaciens Paillot. (Ann. Epiphyt., 8, 1922, 131; L'infection chez les iusectes, 1933, 288.) Larvae of the gypsy moth (Lymantria dispar) are immune to this bacillus. Bacillus bombycoides Paillot. (Compt. rend. Acad. Agr. France, 28, 1942, 158.) Causes lesions because of a bacterial toxin. From infected silkworms. Bacillus bombysepticus Hartman. (Lingnan Sci. Jour., 10, 1931, 280.) Causes a disease of the silkworm (Bom- byx mori). Bacillus borborokoites ZoBell and Up- ham. (Bull. Scripps Inst, of Oceanog- raphy, Univ. Calif., 5, 1944, 274.) Cen- tral spores. From marine bottom deposits. Bacillus borstelensis Stiihrk. (Cent, f. Bakt., II Abt., 93, 1935, 179.) Grows well on Ca n-butyrate agar. Resembles Bacillus rufescens of the same group except that it does not show the typical brown coloration of media. Two strains isolated from soils in German3^ Bacillus bredemannii Chester. (Ba- cillus adhaerens Stiihrk, Cent. f. Bakt., II Abt., 93, 1935, 183; not Bacillus adhaerens Laubach, Jour. Bact., 1, 1916, 503; Chester, in Manual, 5th ed., 1939, 675.) Weak growth on Ca n-butj-rate agar. One strain isolated from Cuban soil. Bacillus bronchitidis Migula. (Ba- cillus der putriden Bronchitis, Lumnit- zer, Cent. f. Bakt., 3, 1888, 621 ; Bacillus bronchitidis putridae Lumnitzer, Wien. med. Presse, 1888, 666; Migula, Syst. d. Bakt., 2, 1900, 641 ; Bacterium lumnitzeri Chester, Man. Determ. Bact., 1901, 120.) 740 MANUAL OF DETERMINATIVE BACTERIOLOGY From sputum in cases of putrid bron- chitis. Bacillus bruneus Migula. (Maschek, Bakt. Untersuch. d. Leitmeritzer Trink- wasser, 1887; Migula, Syst. d. Bakt., 2, 1900, 835; not Bacillus brunneus Eisen- berg, Bakt. Diag., 3 Aufl., 1891, 142.) From water. Bacillus brunneus Eisenberg. (Brauner Wasserbacillus, Adametz and Wich- mann, Die Bakt. d. Nutz- u. Trink- wasser, Mitth. Oesterreich. Ver- suchssta. f. Brauerei u. Mjilzerei, Wien, Heft 1, 1888, 51; Eisenberg, Bakt. Diag., 3 Aufl., 1891, 142; not Bacillus brunneus Schroeter, in Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 158; Bacterium brunneurn Migula, Syst. d. Bakt., 2, 1900, 331.) From water. Non-motile. Bacillus butlerovii Serbinow. (Vest. Russ. obisc. pcelovod. (Messager de la Soc. russe d'Apiculture), No. 3 and No. 11, 1912; see Rev. Appl. Entomol., Ser. A, 1, 1913, 94 and 441.) From black brood of bees. Bacillus butschlii Schaudinn. (Arch, f. Protistenkunde, 1, 1902, 306.) Char- acterized by its large size (3.0 to 6.0 by 24.0 to 80.0 microns) and granular proto- plasm. From the intestine of a cock- roach {Blatta {P eriplaneta) orientalis). Bacillus bulyricus Hueppe. (Hueppe, Mitteil. kaiserl. Gesundheitsamte, 2, 1884, 309; not Bacillus bulyricus Mace, Traits de Bact., 1st ed., 1888; not Bacil- lus bulyricus Botkin, Ztschr. f. Hyg., 11, 1892, 421; Clostridium. Mieppci Trevisan, I generi e le specie delle Batteriacee, 1889, 22; Bacillus pseudobufyricus Kruse, in Flugge, Die Mikroorganismen, 3 Aufl., 2, 1896, 207; Bacillus hueppci Chester, Man. Determ. Bact., 1901, 276.) Bacillus calf actor Miehe. (Arb. der deutsch. Landw. Gesel., Berlin, Heft 3, 1905, 76; Die Selbsterhitzung des Heues, Jena, 1907, 49.) Thought to be the most important thermogenic microorganism in the fermentation of hay. From heating hay. Bacillus canaliculatus Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 20.) From meat extract. Bacillus canceris Migula. (Syst. d. Bakt., 2, 1900, 625.) From a case of stomach cancer. Bacillus caniperda Migula. (Oval- bacillus der Hundestaupe, Galli-Valerio, Cent. f. Bakt., I Abt., 19, 1896, 694; Migula, Syst. d. Bakt., 2, 1900, 764; Bacterium canis Chester, Man. Determ. Bact., 1901, 198.) From nasal mucus and urine of dogs. Bacillus capillaceus Wright. (Mem. Nat. Acad. Sci., 7, 1895, 456.) From water. Bacillus capsici Pavarino and Turconi. (Atti Istit. Bot. R. Univ. Pavia, 15, 1918, 207.) Causes leaf wilt of pepper {Capsicum annuum). May be identical with disease caused by Pseudomonas vesicaloria (Stapp, in Sorauer, Handbuch der Pflanzen-Krankheiten, 2, 5 Aufl., 1928, 262). Bacillus carniphilus Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 19.) From meat extract. Bacillus carnosus Zimmermann. (Tils, Bakt. Untersuch. d. Freiburg. Leitungswasser, Leipzig, 1890, No. 57; Zimmermann, Bakt. unserer Trink- u. NutzwJisser, Chemnitz, II Reihe, 1894, 4.) From water. Bacillus catenulatus Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 27.) Growth on media containing m/100 phenol. Four strains isolated from soil. Bacillus cepae Bassalik and Edelsztein- Kosowa. (Acta Soc. Bot. Polon., 10, 1933, 519.) From diseased onions {Al- lium cepa). Bacillus cerealium Gentner. (Cent. f. Bakt., II Abt., 50, 1920, 428; Pseudo- monas cerealia Stapp, in Sorauer, Handbuch der Pflanzen-Krankheiten, 2, 1928, 22; Bacterium cerealinum Elliott, Manual Bacterial Plant Pathogens, 1930, 111.) Pathogenic for barley {Hordeum vulgar e), rye {Secale cereale) and wheat ( Triticum sp.). Bacillus cinctus Ravenel . (Mem . Nat . Acad. Sci., 8, 1896, 30.) From soil. FAMILY BACILLACEAE 741 Bacillus cirroflagellosus ZoBell and Upham. (Bull. Scripps Inst, of Oceanog- raphy, Univ. Calif., 5, 1944, 266.) Cen- tral spores. Found in marine mud. Bacillus cladoi Trevisan. (Bacille pe- duncul^, Clado, Bull. Soc. Anat. Paris, 1887, 339; Trevisan, I generi e le specie delle Batteriacee, 1889, 14; Bacillus pendunculatus (sic) Eisenberg, Bakt. Diag., 3 Aufi., 1891, 340; Bacillus sepiicus vesicae Sternberg, Man. of Bact., 1893, 475.) From urine in a case of cystitis. Bacillus closteroides Gray and Thorn- ton. (Cent. f. Bakt., II Abt., 73, 1928, 93.) Decomposes phenol. Probably identical with or a variety of Bacillus circidans. Sixteen strains isolated from Rothamsted soils. Bacillus coccineus Pansini. (Pansini, Arch. f. path. Anat., 122, 1890, 437; not Bacillus coccineus Catiano, in Cohn, Beitr. z. Biol. d. Pflanzen, 7, 1896, 339.) From sputum. Red pigment. Bacillus colorans Libermann. (Jour, of Microbiol., Ukraine, 5, 1938, 73; abst. in Cent. f. Bakt., II Abt., 101, 1940, 81.) From fruit conserves containing 10 to 20 per cent sugar. Bacillus comesii Rossi. (Ann. d. Scuola d. Agricult. in Portici, 1903; Arch, di Farmacologia sperim., 3, 1904, fasc. 10.) Similar to Bacillus mesen- tericus. Said to have the abilitj' to dissolve plant particles. Bacillus concoctans Patrick and Werk- man. (Iowa State Coll. Jour. Sci., 7, 1933, 415.) Ferments xylan. One cul- ture isolated from soil. Bacillus conjunctivitidis subtiliformis Michalski. (Cent. f. Bakt., I Abt., Orig., 36, 1904, 212.) From more than 50 cases of acute conjunctivitis. Similar to Bacillus subtilis. Bacillus consolidus Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 191.) Good growth on Ca n-butyrate agar. One strain isolated from Cuban soil. Bacillus contextus Migula. {Bacillus D, Peters, Botan. Zeitung, 47, 1889; Migula, Syst. d. Bakt., 2, 1900, 522.) From leaven. Bacillus corrugatus Migula. {Bacillus viesentericus vulgalus Fliigge, Die Mikro- organismen, 2 Aufl., 1886, 322; Bacillus Xo. II, Fliigge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lactis No. II, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 208; iMigula, Syst. d. Bakt., 2, 1900, 583.) From milk. Bacillus corruscans Schroeter. (In Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886 r 158.) From cooked potato. Bacillus costatus Lloyd. (Jour. Bact., 21, 1931, 94.) From sea water off Scot- land. Nitrates and nitrites reduced to nitrogen. Bacillus crinatus Chester. (Bacillus No. 5, Pansini, Arch. f. path. Anat., 122, 1890, 441; Chester, Man. Determ. Bact., 1901, 281.) From sputum. Bacillus crinitus Wright. (Wright, Mem. Nat. Acad. Sci., 7, 1895, 453; Bacterium crinatum (sic) Chester, Man. Determ. Bact., 1901, 192.) From river water. Bacillus crystalloides Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 371; Bacterium crystaloides (sic) Chester, Man. Determ. Bact., 1901, 191.) From the air. Bacillus cubensis Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 192.) Good growth on Ca n-butyrate agar. Two cultures isolated from soils from Cuba. Bacillus cystiformis Trevisan. (Ba- cille cystiforme, Clado, Bull. Soc. Anat. Paris, 1887, 339; Trevisan, I generi e le specie delle Batteriacee, 1889, 14.) From urine in a case of cystitis. Bacillus cytaseus McBeth and Scales. (U. S. Dept. Agr., Bur. Plant Industry, Bull. 266, 1913, 39; Bacterium cytaseum Holland, Jour. Bact., 5, 1920, 218.) Requires cellulose for best growth. From decomposing materials and soil. Bacillus cytaseus var. zonalis Keller- man et al. (Cent. f. Bakt., II Abt., 39, 1913,511.) From soil from Utah. While no spores were observed, this organism was like Bacillus cytaseus except that colonies on cellulose agar showed con- 742 MANUAL OF DETERMINATIVE BACTERIOLOGY eentric opaque or semi-opaque and transparent zones. Bacillus danteci Kruse. (Bacille rouge de Terre-Neuve, Le Dantec, Ann. Inst. Past., 5, 1891, 662; Kruse, in Flugge, Die Mikroorganismen, 3 Aufl., 2, 1896, 270.) From reddened salt cod fish. Bacillus daucarum von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 494.) Appar- ently a strain of Bacillus subtilis. From canned carrots. Bacillus demmei Trevisan. (Bacillus der Erythema nodosum, Demme, Fort- schr. d. Med., 6, 1888, 257; Trevisan, I generi e le specie delle Batteriacee, 1889, 14 ; Bacillus erythematis Kruse, in Fliigge, Die ]Mikroorganismen, 3 Aufl., 2, 1896, 426; Bacillus erythematis maligni Kruse, ihid., 479; Bacterium erythematis Migula, Syst. d. Bakt., 2, 1900, 346.) From erythema nodosum (skin). Bacillus dendroides Holzmtiller. (Cent. f. Bakt., II Abt., 23, 1909, 331.) From frog feces. Closely related to Bacillus mxjcoides. Bacillus dendroides Thornton. (Thornton, Ann. Appl. Biol., 9, 1922, 247; not Bacillus dendroides Holzmiiller, loc. cit.) Common in Rothamsted soil. Said to belong to the Bacillus subtilis group. Bacillus dentatus Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 106.) Good growth on valine agar. Two cultures iso- lated from soil of Jugoslavia and North Carolina. Bacillus destruens von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 502.) From boiled asparagus. Bacillus detrudens Wright . (Mem . Nat . Acad. Sci., 7, 1895, 452.) From water. Bacillus diastalicus Boyarska. (Iz- vestia Acad. Sci., U. S. S. R., Biol. Ser., 1941.) Thermophilic. Bacillus disciformans Zimmermann. (Zimmermann, Bakt. unserer Trink. u. Nutzwasser, Chemnitz, II Reihe, 1894, 48; Bacterium disciformans Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 238.) From waste water. Appar- ently not identical with Bacillus disci- formis Grafenhan, although the name suggests possible relationship. Bacillus disciformis Grafenhan. (In- aug. Diss., Halle, 1891, 1.) From water. From the description, this organism may be Bacillus subtilis. Bacillus distortus (Duclaux) Trevisan. {Tyrothrix distortus Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From milk. Bacillus dobelli Duboscq and Grass^. (Arch. Zool. Exper. et G^n., 66, 1927, 487; Bacillus (Flexilis) dobelli Duboscq and Grassd, ibid., 487.) Similar to Ba- cillus flexilis Dobell. Found in rectum of a termite {Calotermes (Glyptotermes) iridipennis). These authors suggest that Bacillus flexilis Dobell, Bacillus bulschlii Schaudinn and Bacillus dobelli be grouped under the name Flexilis. Bacillus duclauxii (Miquel) Chester. (Bacillus ureae Miquel, Bull. Soc. Chim. d. Paris, 31, 1879, 391; Urobacillus du- clauxii sive Bacillus ureae /3 Miquel, Ann. d. ]\Iicrog., 2, 1889-1890, 53, 122 and 145; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 10, 1898, 123.) From water and soil. Bacillus dysodes Zopf. (Die Spalt- pilze, 3 Aufl., 1885,' 90.) From ferment- ing dough. Bacillus elegans Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 103.) Four cultures isolated from soil, one from Jugoslavia and three from Germany. Bacillus emulsionis Beijerinck. (Folia Microbiol., 1, 1912, 377; see Perquin, Jour. Microbiol, and Serol., 6, 1940, 226.) Produces slime in sucrose solutions. Bacillus encephaloides Trevisan. (Ba- cille de I'air /, Babes, in Cornil and Babes, Les Bacteries, 1886, 150 ; Trevisan, I generi e le specie delle Batteriacee, 1889,20.) From the air. Bacillus enterothrix Collin. (Arch. Zool. Exp(5r. et G^n., 51, 1913, Notes and Revue, No. 3.) Found in the rectum of toad tadpoles (Alytes sp.). Bacillus epidermidis (Bizzozero) Bor- doni-Uffreduzzi. (Leptothrix epidermi- FAMILY BACILLACEAE 743 dis Bizzozero, Arch. f. path. Anat., 98, 1884, 441 ; Bordoni-Uffreduzzi, Fortschr. d. Med., 4, 1886, 156; Carcinombacillus, Scheuiien, Deutsche med. Wochiischr., 1887, 1083; Bacillus mesentericus ruhigi- nosus Senger, Cent. f. Bakt., 3, 1888, 603; Bacillus bizzozerianus Trevisan, I generi e le specie delle Batteriacee, 1889, 14; Bacillus scheurleni Dyar, Ann. X. Y. Acad. Sci., 8, 1895, 367.) From the human mouth and skin. Mace (Traite pratique de Bact., 4th ed., 1901, 1071) says that this organism is the ordinary' potato hacillus, i.e., Bacillus I'ulgatus. Bacillus epiphytus ZoBell and Upham. fBuU. Scripps Inst, of Oceanography, Cniv. Calif., 5, 1944, 266.) Central spores. Found associated with marine phj'toplankton. Bacillus erodens Ravenel. (^lem. Xat. Acad. Sci., 8, 1896, 35.) From soil. Bacillus eslerificans ]\Iaassen. (Arb. a. d. kaiserl. Gesundheitsamte, 15, 1899, 504; Plectridium esterificans Huss, Cent, f. Bakt., II Abt., 19, 1907, 52.) Found in a solution of decomposing litmus; also isolated from butter. Bacillus evanidus Grohmana. (Mor- phologisch-biologische Beitrage zur Kenntnis der Wasserstoffbakterien, In- aug. Diss., Univ. Leipzig, 1923; Cent. f. Bakt., II Abt., 61, 1924, 207; Ruhland and Grohmann, Jahrb. Wissensch. Bota- nik, 63, 1924, 321.) Oxidizes hydrogen in the presence of oxygen to form water. Presumably widely distributed in soil. Bacillus exiguus Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 44.) Isolated 3 times from gar- den air. Bacillus exilis Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 29.) Growth in media containing m/100 phenol. Eight strains isolated from soil. Bacillus fastidiosus den Dooren de Jong. (Cent. f. Bakt., II Abt., 79, 1929, 344.) Six strains isolated from unheated garden soil. Bacillus ferrigenus Bargaglio-Petrucci. (Xuovo Giornale botanico italiano, 1913, 1914, 1915; quoted from De Rossi, Microbiol. Agraria e Technica, 1927, 904.) A facultative thermophile, grow- ing up to 65° to 70°C. Bacillus feslin^is McBeth. (Soil Sci., 1, 1916, 451.) Filter paper decomposed to a grayish-white felt-like mass. From soil in California. Bacillus filamentosus Klein. (Klein, see Migula, Syst. d. Bakt., 2, 1900, 285; Bacterium filamentosum Burchard, Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1902, 22.) Bacillus filaris ^ligula. (Bacillus Xo. XI, Flligge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus lactis Xo. XI, Kruse, in Fli'igge, Die Mikroorganismen, 3 Aufl., 2, 1896, 209 ; :\Iigula, Syst. d. Bakt., 2, 1900, 579; Bacillus aromaiicus Chester, Man. Determ. Bact., 1901, 276; not Bacillus aromaticus Pammell, Bull. Xo. 21, Iowa Agr. Exp. Sta., 1893, 792; not Bacillus aromaticus Grimm, Cent. f. Bakt., II Abt., 8, 1902, 584.) From milk. Bacillus filicolonicus ZoBell and Up- ham. (Bull. Scripps Inst, of Oceanog- raphy, Univ. Calif., 5, 1944, 270.) Sub- terminal spores. From sea water and marine mud. Bacillus filiformis (Duclaux) Trevisan. {Tyrothrix filiformis Duclaux, Ann. Inst. Xat. Agron., 4, 1882, 23; Trevisan, I generi e le specie delle Batteriacee, 1889, 16; not Bacillus filiformis Tils, Ztschr. f. Hyg., 9, 1890, 317; not Bacillus filiformis Migula, Syst. d. Bakt., 2, 1900, 587.) From cheese. Bacillus fissuratus Ravenel. (]\Iem. Xat. Acad. Sci., 8, 1896, 38.) From soil. Bacillus fitzianus Zopf. (Fitz, Ber. d. deutsch. chem. Gesellsch., 6, 1873, 48; ibid., 9, 1876, 1348; ibid., 10, 1877, 276; Glycerinaethylbacterie, Buchner, in Xa- geli, Untersuch. (i. niedere Pilze, 1882, 220 ; Zopf, Die Spaltpilze, 1 Aufl., 1883, 52 ; Bacterium fitzianum Zopf, Die Spaltpilze, 2 Aufl., 1884, 49.) From boiled hay infusions. Forms butyric acid. Bacillus flagellifer Migula. (Bacillus Xo. VI, Flligge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lactis Xo. VI, Kruse, in Flligge, Die Mikroorganismen, 3 Aufl., 2, 1896, 209; Migula, Syst. d. Bakt., 744 MANUAL OF DETERMINATIVE BACTERIOLOGY 2, 1900, 581 ; Bacillus rudis Chester, Man. Determ. Bact., 1901, 279.) From milk. Bacillus flavescens Weiss. (Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 258; not Bacillus flavescens Pohl, Cent. f. Bakt., 11, 1892, 144.) From brewer's grains. Uncommon. Bacillus flavidus Sttihrk. (Cent. f. Bakt., II Abt., 93, 1935, 185; not Bacillus flavidus Fawcett, Rev. Indust. y Agrico. de Tucuman, 13, 1922, 5; not Bacillus flavidus Morse, Jour. Inf. Dis., 11, 1912, 284.) Good growth on Ca n-butyrate agar. One culture isolated from soil from Egypt. Bacillus flavidus aZreiKlamann. (Bien- enwirtschaftl. Cent., Hanover, 1890, No. 2.) Associated with foulbrood of bees. Bacillus flavoviridis Migula. (Mas- chek, Bakt. Untersuch. d. Leitmeritzer Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 821.) From water. Bacillus flexilis Dobell. (Quart. Jour. Microsc. Sci., 52, 1908, 121; Arch. f. Protistenk., 26, 1912, 117.) Reported as being similar to Bacillus butschlii Schau- dinn although only half its size. From the large intestine of frogs (Rana tem- poraria) and toads {Bufo vulgaris). Bacillus flexus Batchelor. (Jour. Bact., 4, 1919, 23.) Resembles Bacillus megatherium. From intestinal contents of a child. Bacillus fiuorescens undulalus Ravenel. (Mem. Nat. Acad. Sci., 8, 1S96, 20.) From soil. Bacillus foliaceus Migula. {Bacillus mesentericus fuscus Fltigge, Die Mikro- organismen, 2 Aufl., 1886, 321; Bacillus No. IV, Flugge, Ztschr. f. Hyg., 17, 1894, 294; Bacillus lactis No. IV, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 208; Migula, Syst. d. Bakt., 2, 1900, 582.) From milk, air and soil. Bacillus for mosus Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 101 ; not Bacillus formosus Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 12.) One culture isolated from soil from Montenegro. Bacillus foutini Chester. (Bacillus D, Foutin, Cent. f. Bakt., 7, 1890, 373; Chester, Man. Determ. Bact., 1901, 285.) From hail. Bacillus frankei (sic) DeToni and Trevisan. (Sarkombacillen, Francke, Munch, med. Wochnschr., 1888, No. 4; abst. in Cent. f. Bakt., 3, 1888, 601; DeToni and Trevisan, in Saccardo, Syl- loge Fungorum, 8, 1889, 967. ) From cases of sarcoma. Bacillus freudenreichii (Miquel) Ches- ter. (Urobacillus freudenreichii sive Ba- cillus ureae y Miquel, Ann. d. Micro- graphie, 2, 1890, 367 and 488; Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 10, 1898, 110.) Lohnis (Cent. f. Bakt., II Abt., U, 1905, 719) considered this a variety of Bacillus pumilus. Gibson (Jour. Bact., 29, 1935, 493) believed it belonged to the Bacillus pasteurii group although no authentic cultures were available. From soil. Bacillus frutodestruens Madhok and Ud-Din. (Indian Jour. Agr. Sci., 13, 1943, 129.) Causes a rot of tomato fruit. Bacillus funicularis Kluyver and Van Niel. (Planta, Arch. f. wissensch. Bota- nik, 2, 1926, 507.) Exhibits contact irritability. From soil. Bacillus furvus Goadby. (Dental Cosmos, Jt2, 1900, 322.) From the mouth. .Vssociated with dental caries. Bacillus (Streptobacter) fusisporns Schroeter. (In Cohn, Krypt. Flora v. Schlesien, 3, 1, 1886, 161.) From waste water in a sugar factory. Bacillus fusus Batchelor. (Jour. Bact., 4, 1919, 27.) Said to resemble Bacillus centrosporus, i.e., Bacillus brevis. From feces. Bacillus gangliformis Ravenel. (Rav- enel, Mem. Nat. Acad. Sci., 8, 1896, 34; Bacterium gangliforme Chester, Man. Determ. Bact., 1901, 193.) From soil. Bacillus gangraenae Chester. {Bacil- lus gangraenae pulpae Arkovy, Cent. f. Bakt., 23, 1897, 921; Chester, Man. Determ. Bact., 1901, 275; not Bacillus gangraenae Tilanus, Nederl. Tijdschr. v. FAMILY BACILLACEAE 745 Geneeskunde, 21 , 1885, 110.) Associated with gangrene of tooth pulp and caries of teeth. Bacillus gasofortnans Pammel. (Pani- mel, Cent. f. Bakt., II Abt., 2, 1896, &42; not Bacillus gasofortnans Eisenberg, Bakt. Diag., 3 Aufi., 1891, 107; Bacillus pajnmelii Chester, Man. Determ. Bact., 1901,270.) From cheese. Bacillus gelalinosus ^Nligula. {Bac- ieriurn gelatinosutn betae Glaser, Cent. f. Bakt., II Abt., /, 1895, 879 ; Migula, Syst. d. Bakt., 2, 1900, 805.) Produces slime in sucrose solutions. Probably a variety of Bacillus vulgalus according to Sac- chetti (Cent. f. Bakt., II Abt., 95, 1936, 115). Bacillus geniculatus (Duclaux) Trevi- san. (Tyrothrix geniculatus Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 16; not Bacillus genicula- tus De Bary, Inaug. Diss., Strassburg, Leipzig, 1885; not Bacillus geniculatus Wright, Mem. Nat. Acad. Sci., 7, 1894, 459; Bacillus gonatodes DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 964; Bacterium geniculatum Migula, Syst. d. Bakt., 2, 1900, 322.) From milk. Bacillus gigas (Koch) Trevisan. {Streptobacteria gigas-pericardii Billroth, Die Vegetationsformen von Coccobacteria septica, Berlin, 1874, 60; Streptobacteria gigas Koch, in Cohn, Beitr. z. Biol. d. Pflanz., 2, Heft 3, 1877, 429; Metallacter gigas Trevisan, Batter, ital., 1879, 25; Trevisan, Atti. d. Accad. Fis.-Med.- Stat., Milan, Ser. 4, 3, 1886, 96; not Bacillis gigas van der Goot, Med. Proef- station voor de Java Suikerindust., pt. 5, No. 10, 1915, 284; not Bacillus gigas Zeissler and Rassefeld, Arch. f. wiss. u. prakt. Tierheilk., 59, 1929, 419.) From pericardial exudate. Bacillus ginglymus Ravenel. (Mem. Nat. Acad. Sci., 8, 1896, 37.) From soil. Bacillus glaciformis Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 29.) From meat extract. Bacillus globifer Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 26.) Growth on media containing m/100 phenol. Author considers it similar to Bacillus alvei. Five strains isolated from soil. Bacillus gluiinis Patrick and Werk- raan. (Iowa State Coll. Jour. Sci., 7, 1933, 414.) Ferments xylan. Two strains isolated from decayed apple wood. Bacillus glycinophilus Rippel. (Rip- pel, Arch. f. :Mikrobiol., 8, 1937, 42; also see Rippel andNabel, ibid., 10, 1938,359.) From intestines of cattle. Fresh cultures on agar form protein from glycine and glucose. Bacillus gussypina Stedman. (Ala- bama Agr. Exp. Sta. Bui. 55, 1894, 6; Earle, Alabama Agr. Exp. Sta. Bui. 107, 1899, 311.) Reported as cause of boll rot on cotton {Gossypium sp.) Bacillus granularis Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 180.) Very good growth on Ca n-butyrate agar. One culture isolated from garden soil in Germany. Bacillus granulosus Russell. (Russell, Ztschr. f. Hyg., 11, 1892, 194; Bacterium granulosum Chester, Man. Determ. Bact., 1901, 194.) From mud from the Gulf of Naples. Bacillus grossus Migula. (Bacteri- enart No. 13, Lembke, Arch. f. Hyg., 26, 1896, 308; Migula, Syst. d. Bakt., 2, 1900, 570.) From the intestines of infants. Bacillus gryllotalpae Metalnikov and Meng. (Compt. rend. Acad. Sci., Paris, 201, 1935, 367.) From diseased larvae of the cricket {Gnjllotalpa gryllotalpa). Bacillus guano Stapp. (Cent. f. Bakt., II Abt., 51, 1920, 29.) From soil pre- viously fertilized with guano. Bacillus gummosus Happ. (Bakt. und Chem. Untersuch. iiber die schleimige Gahrung. Univ. Basel, 1893, 34; abst. in Cent. f. Bakt., U, 1893, 175.) From digitalis infusions. Presumably a mu- coid form of a common spore-former. See Bacterium gummosujn Ritsert. Bacillus harai Hori and Mij^ake. (Rpt. Imp. Agr. Exp. Sta. Nishigahara, 38, 746 MANUAL OF DETERMINATIVE BACTERIOLOGY 1911, 69.) Pathogenic for willow {Salix sp.). Bacillus hessii (Guillebeau) Kruse. {Bacterium hessii Guillebeau, Landw. Jahrb. d. Schweiz, 5, 1891, 138; Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 210.) There is some question whether the original culture was a spore- former or whether it was mixed with one of the common slimy milk organisms. From slimy milk. Bacillus iiirudinis Schweizer. (Arch, f. Mikrobiol., 7, 1936, 235.) From the digestive slime of leeches (Hirudo medi- cinalis and Hirudo officinalis) . Bacillus hollandicus Stapp. (Cent. f. Bakt., II Abt., 51, 1920, 47.) From soil from Delft. Bacillus hoplosternus Paillot. (Compt. rend. Acad. Sci., Paris, 163, 1916, 774; Ann. Inst. Past., S3, 1919,403.) Isolated once from the body fluid of a June bug. Pathogenic for several species of insects. Bacilhis iviminutus McBeth. (Soil Sci., 1, 1916, 455.) Growth only in the presence of cellulose. From ten different soils of California. Bacillus immohilis Steinhaus. (Jour. Bact., Jt2, 1941, 783.) The author states that it probably belongs to the Bacillus adhaerens group. From rectum of larvae of the sphinx moth {Ceratomia catalpae). Bacillus iniomarinus ZoBell and Up- ham. (Bull. Scripps Inst, of Oceanog- raphy, Univ. Calif., 5, 1944, 265.) Sub- terminal spores. From marine bottom deposits in shoal waters. Bacillus indifferens Soriano. (Thesis, Univ. Buenos Aires, 1935, 565.) One strain isolated from soil . Bacillus infantilis Kendall. (Jour. Biol. Chem., 5, 1909, 419 and 439.) From the intestine in cases of infantilism. Saprophj^tic. Bacillus intermittens Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 23.) From meat extract. Bacillus intrapallans Forbes. (Bull. Illinois State Lab. Natur. Hist., Art. IV, 1886, 283, 288 and 297.) Bacillus jubatus Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 24.) Very good growth on media containing m/50 phenol. Nine strains isolated from soil. Bacillus kaleidoscopicus Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 31.) From meat extract. Bacillus kefir Kuntze. (Cent. f. Bakt., II Abt., U, 1909, 116.) From kefir, a Caucasian milk beverage. Bacillus kermesinus Migula. (Kar- minroter Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 67; Migula, Syst. d. Bakt., 2, 1900, 858.) From water. Bacillus kildini Issatchenko. (Re- cherchcs sur les Microbes de I'Ocdan Glacial Arctique. Petrograd, 1914.) From sea water. Bacillus koubassoffii Chester. (Bacil- lus der krebsartigen Neubildungen, Koubassoff, Vortrag. Moskauer Mili- tiirarztlichen Verein, 1888, No. 22; abst. in Cent. f. Bakt., 7, 1890, 317; Chester, Man. Determ. Bact., 1901, 282.) From cancerous growths of the human stomach. Bacillus lacca Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 411.) From the stomachs and intestines of birds. Bacillus lactis-albus Chester. {Bacil- lus lactis albus Sternberg, Man. of Bact., 1893, 680; Chester, Man. Determ. Bact., 1901,277.) From milk. Bacillus lactucae Voglino. (Ann. R. Accad. Agr. Torino, 4(>, 1903, 25.) Patho- genic for lettuce {Lactuca saliva). Bacillus lasiocampa Brown. (Amer. Museum Novitates, No. 251, 1927, 7.) Said to belong to Bacillus subtilis group. From ovaries and egg tubes of tent cater- pillar moth. Bacillus latvianus Kalnins. (Latvijas Univ. Raksti, Serj^a I, No. 11, 1930, 265.) Cellulose attacked at 34°C but not at 37°C. Probably from soil. Bacillus lautus Batchelor. (Jour. Bact., //, 1919, 30.) One culture from feces. Bacillus legrosii Hauduroy et al. (Legros, These Med. Paris, 1902; Haudu- roy et al., Diet. d. Bact. Path., 1937, 43.) FAMILY BACILLACEAE 747 Facultative anaerobe producing gaseous gangrene. From a gaseous suppuration. Bacillus legumini-perdus von Oven. (Cent. f. Bakt., II Abt., 16, 1906, 67; Bacteriian leguminiperdum Stevens, The Fungi which Cause Plant Disease, 1913, 28.) Pathogenic for lupine {Lupinus sp.), kidney bean {Phaseolus vulgaris), pea {Pisuni sativu77i) , tomato {Lycopersi- cimi esculenium) . Bacillus lehmanni Herter. (Microbe 5A, Choukevitch, Ann. Inst. Past., 25, 1911, 350; Herter, Just's Botan. Jahres- ber., 3.9, 2 Abt., Heft 4, 1915, 748.) From the large intestine of the horse. Bacillus lekitosis Perlman. (X. Y. State Dept. Agr. and Markets, Ann. Rept., 1931, 115.) Egg lecjjthin hydro- lyzed completely by powerful extracellu- lar enzyme. From contaminated sample of whole mixed eggs. Bacillus lesagei Trevisan. (Bacille de la diarrhee verte des enfants, Lesage, Bull. Acad. Med. Paris, Oct. 25, 1887; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 14.) From the intestine of infants. Bacillus levanijormans Greig Smith. (Proc. Linn. Soc. New South Wales, 26, 1901, 589, 674 and 684; Cent. f. Bakt., II Abt., 8, 1902, 596.) Produces slime in sucrose solutions. Probably a variety of Bacillus vulgatus according to Sacchetti (Cent. f. Bakt., II Abt., 95, 1936, 115). Bacillus licheniformis (Weigmann) Chester. (Bacterie a, Weigmann, Cent, f . Bakt., II Abt., 2, 1896, 155 ; Clostridium licheniforme Weigmann, loc. cit., 4, 1898, 822; Chester, Man. Determ. Bact., 1901, 287; see also Gibson, Soc. Agric. Bact. (British); Abstr. Proc, 1927, Paper Xo. 10; Gibson and Topping, ibid., 1938, 43; Gibson, ibid., 1943, 13.) Gibson places this with Bacillus subtilis although it was originally described as being Gram- negative and forming clostridial spor- angia. Spore germination polar. Bacillus lichenoides Grohmann. (Mor- phologisch-biologische Beitrage zur Kenntnis der Wasserstoffbakterien, In- aug. Diss., Univ. Leipzig, 1923; Cent. f. Bakt., II Abt., 61, 1924, 267 ; Ruhland and Grohmann, Jahrb. wissensch. Botanik, 63, 1924, 321.) Oxidizes hydrogen in the presence of oxygen to form water. Pre- sumably widely distributed in soil. Bacillus lignivorans Patrick and Werk- man. (Iowa State Coll. Jour. Sci., 7, 1933, 410.) Ferments xylan. One cul- ture isolated from decayed maple wood. Bacillus lignorum Patrick and Work- man. (Iowa State Coll. Jour. Sci., 7, 1933, 410.) Ferments xylan. One cul- ture isolated from decayed apple wood. Bacillus limnophilus Stiihrk. (Cent, f. Bakt., II Abt., 93, 1935, 190.) Good growth on Ca n-butyrate agar. One culture isolated from soil of Germany. Bacillus lingardi Trevisan. (Bacillus de la stomatite ulcereuse du veau, Lin- gard and Batt ; Trevisan, I generi e le specie delle Batteriacee, 1889, 14.) From ulcerations on the tongue and mucous membrane of the mouth of calves. Bacillus lividus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 18; not Bacillus lividus Voges, Cent. f. Bakt., 14, 1893, 303.) From water. Bacillus longior Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 57. ) Isolated once from garden air. Bacillus longus Migula. (Bacillus No. VII, Fltigge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus lactis X'o. VII, Kruse, in Fltigge, Die Mikroorganismen, 3 Aufi., 2, 1896, 209; Migula, Syst. d. Bakt., 2, 1900, 581 ; not Bacillus longus Chester, Man. Determ. Bact., 1901, 303; Bacillus plicatus Chester, ibid., 275; not Bacillus plicatus Frankland and Frankland, Philos. Trans. Royal Soc. London, 178, B, 1887, 273; not Bacillus plicatus Zim- mermann, Bakt. unserer Trink- u. Nutzwasser, Chemnitz, I Reihe, 1890, 54; not Bacillus plicatus Deetjen, Inaug. Diss., Wiirzburg, 1890; not Bacillus plicatus Copeland, Rept. Filtration Com- mission, Pittsburgh, 1899, 348.) From milk. 748 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacillus loxosporus Burchard. (Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1902, 49.) From the air. Synonym of Bacil- lus simplex according to Gottheil, Cent. f. Bakt., II Abt., 7, 1901,633. Bacillus luteus Garbowski. (Bacillus luteus sporogenes Smith and Baker, Cent. f. Bakt., II Abt., 4, 1898, 788; Garbowski, Cent. f. Bakt., II Abt., 19, 1907, 641.) From two samples of beet sugar. Bacillus lutzae Brown. (Amer. Museum Novitates, No. 251, 1927, 8.) Pathogenic for certain flies. Dying indi- viduals of the green blow fly (Lucilia sericata) yielded pure cultures. Bacillus maculatus Sttihrk. (Cent. f. Bakt., II Abt., 93, 1935, 184.) Good growth on Ca n-butyrate agar. Two cultures isolated from soils from Cuba and Germany. Bacillus maidis Paltauf and Heider. (Paltauf and Heider, Wiener med. Jahrb., 3, 1888, 383; Paltauf, Med. Jahrb., No. 8, 1889.) From an infusion of maize; also from feces in cases of pellagra. This species was originally described by Cuboni, probably in the Rendic. R. Accad. dei Lincei, /, 1886. It was later shown to be a spore-former of the Bacil- lus viesentericus group. It was quite different from the organism isolated by Tataroff {Pseudoiywnas maidis Migula) and identified by him as Bacillus mai- dis. Bacillus malakofaciens von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 499.) Reported to be similar in morphology and physiology to Bacillus asterosporus. From preserved asparagus and from green beans. Bacillus maritimus Russell. (Russell, Bot. Gazette, 18, 1893, 445; Bacterium maritimum Chester, Man. Determ. Bart., 1901, 189.) From sea-mud. Bacillus mazun Weigmann, Gruber and Huss. (Cent. f. Bakt., II Abt., 19, 1907, 72.) From the Armenian milk product, mazun. Bacillus mediosponts Migula. (Bacil- lus No. VIII, Fliigge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus laciis No. VIII, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 209; Migula, Syst. d. Bakt., 2, 1900, 580; Bacillus viagnus Chester, Man. Determ. Bact., 1901, 276.) From milk. Bacillus medio-tumescens Saito. (Jour. Coll. Sci. Imp. Univ., Tokyo, 23, Art. 15, 1908, 45.) Isolated twice from garden air. Bacillus megatherium Ravenel. (Ravenel, Mem. Nat. Acad. Sci., 8, 1896, 11; not Bacillus megatherium De Bary, Vergleich. Morph. u. Biol. d. Pilze, 1884, 499; Bacillus megatherium var. ravenelii Chester, Man. Determ. Bact., 1901, 271.) From soil. Bacillus melanosporus Schroeter. (Eine Bacillus Art, Eidam, Beitr. z. Biol. d. Pflanz., 1,3, 1875,216; Schroeter, in Cohn, Kryptog. Flora v. Schlesien, 3, 1, 1886, 159.) From cooked potato. Bacillus melonis Patrick and Werkman. (Iowa State Coll. Jour. Sci., 7, 1933, 411 ; not Bacillus vielonis Giddings, Vermont Agr. E.xp. Station Bull. 148, 1910, 413.) Ferments xylan. One culture isolated from decayed watermelon. Bacillus iuesentericus fuscus consistens Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 373.) Found as a contamination in a milk culture. Bacillus meseniericus fuscus granidatus Dyar. (Ann. N. Y. Acad. Sci., 8, 1895, 373.) Found abundantly in a jar of sterilized milk. Bacillus meseniericus roseus Zimmer- mann. (Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, 2, 1894, 26.) From water. Zimmermann received this cul- ture from Krai under the above name. Krai (Sanmilung v. Mikroorganismen, Prague, 1900, 7) lists it as a synonym of Bacillus meseniericus ruber Globig. Bacillus mesentericus vulgatus rnucosus Ivanovics. (Cent. f. Bakt., I Abt., Orig., 142, 1938, 52.) Author believed it to be identical or near to Bacillus vulgatus FAMILY BACILLACEAE 749 but produces much slime. From drain- age water. Bacillus mesenterioides Deetjen. (In- aug. Diss., Wiirzburg, 1890.) From sausage. Bacillus viicans Wilhelmy. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 30.) From meat extract. Bacillus milii (sic) Howard. (Glean- ings in Bee Culture, 28, 1900, 124.) From black brood of the honey bee (Apzs mellifera) . Bacillus initochrondrialis Alexeieff. (Arch. f. Protist., Jfi, 1924, 399.) From horse manure. Bacillus modestus Schieblich. (Cent, f. Bakt., I Abt., Orig., 124, 1932, 269.) Prefers carbohydrate media and 37°C. From grass and meadow plants. Bacillus monachae (von Tubeuf) Eck- stein. (Bacillus B, Hofmann, Die Schlaffsucht (Flacherie) der Nonne {Liparis monacha), 1891, Frankfort am Main ; Bacterium monachae von Tubeuf, Forstl. naturwissensch. Ztschr., /, 1892, 34, 277; Eckstein, Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 6.) From diseased caterpillars of the nun moth (Lijmantria monacha). Bacillus montanus Werner. (Cent. f. Bakt., II Abt., 87, 1933, 458.) Good growth on Ca n-butyrate agar. One cul- ture isolated from soil of Germany. Bacillus moulei DeToni and Trevisan. (Bacille des viandes a odeur de beurre ranee des halles de Paris, Nocard and Moule, Recueil de m^decine vet^r., Ser. 7, 6, 18 — , 67; DeToni and Trevisan, in Sacardo, Sylloge Fungorum, 8, 1889, 971.) From spoiled meat. Bacillus mucilaginosus Migula. (Happ, Inaug. Diss., Berlin, 1893, 28; Migula, Syst. d. Bakt., 2, 1900, 696.) From a slimy fermentation. Bacillus mucosus Zimmerman. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 8; Bacterium mucosum Migula, Syst. d. Bakt., 2, 1900, 315.) From water. Bacillus mucronatus Saito . (Jour . Coll . Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 58.) Isolated twice from garden air. Bacillus multipediculus flavus Zimmer- mann. (Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, II Reihe, 1894, 42.) From water. Bacillus muralis Tomaschek. (Botan. Zeit., 46, 1887,665.) Bacillus mutabilis Soriano. (Estudio sistematico de algunas bacterias esporu- ladas aerobias. Thesis, Univ. Buenos Aires, 1935, 570.) Four strains isolated from fecal matter and Argentine soils. Bacillus myxodens Burchard. (Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1898, 41.) From beer yeast. Bacillus nebulosus Goresline. (Gores- line, Jour. Bact., ^7, 1934, 72 ; not Bacillus nebulosus Wright, Mem. Nat. Acad. Sci., 7, 1894, 465 ; not Bacillus nebulosus Halle, These de Paris, 1898; not Bacillus nebu- losus Vincent, Ann. Inst. Past., 21, 1907, 69 ; not Bacillus nebulosus gazogenes Jacobson, Ann. Inst. Past., 22, 1908, 300.) From a trickling filter receiving creamery wastes. Bacillus neumanni Herter. (Microbe 5B, Choukevitch, Ann. Inst. Past., 25, 1911, 350; Herter, Just's Botan. Jahres- ber., 39, 2 Abt., Heft 4, 1915, 748.) From the large intestine of the horse. Bacillus nigrescens Bartels. (Cent. f. Bakt., II Abt., 103, 1940, 22.) Good growth on media containing m/25 phenol. Three strains isolated from soil. Bacillus nigricans Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 428.) From the stomach of a bird. Bacillus nitidis Heigener. (Cent. f. Bakt., II Abt., 53, 1935, 99.) One culture isolated from soil from Washington, D. C. Bacillus nitri Ambroz. (Cent. f. Bakt., I Abt., Orig., 51, 1909, 193.) Used for cytological studies because of its large size. Found on gelatin plates poured for the isolation of yeasts. Bacillus nitroxus Beijerinck. (Cent, f. Bakt., II Abt., 25, 1909, 45.) In the absence of air forms N gas, CO2, and N2O in nitrate broth. Under aerobic condi- 750 MANUAL OF DETERMINATIVE BACTERIOLOGY tions only a weak reduction. From garden soil. Bacillus nobilis Adametz. (See Freud- enreich, Cent. f. Bakt., II Abt., 7, 1901, 857; ibid., 8, 1902, 674.) This organism was sold under the name Tyrogen; it was said to play a part in the ripening of hard cheese. This was doubted by Freudenreich who identified it as one of the Tyrothrix group. Original descrip- tion apparently in Osterreichen Mokerei- Zeitung, Nov. 15, 1900; Dec. 1 and 15, 1900; Milchzeitung, No. 4S, 1900. Bacillus novus (Huss) Bergey et al. {Plectridium novum Huss, Cent, f . Bakt., II Abt., 19, 1907, 256; Bergey et al., Manual, 1st ed., 1923, 304.) From steril- ized milk. Bacillus oblongus Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 16.) From the larvae of a moth {Htjpono- meuta cvonymella) . Bacillus oehensis Bartels. (Cent. f. Bakt., II Abt., 108, 1940,28.) Growthon media containing m/100 phenol. One culture isolated from soil. Bacillus oleae Schiff-Giorgini. (Cent. f.Bakt., II Abt., 15, 1905,203.) Thought to be the cause of tubercles on the olive tree from which it was isolated. Bacillus omelianskii Serbinoff. (Zhur- nal Bolezni Rastenii, Leningrad, 9, 1915, 105.) Causes a rot of sorghum. Bacillus ontarioni (Chorine) Stein- haus. {Bacterium ontarioni Chorine, Internat. Corn Borer Invest., Sci. Rpts., 2, 1929, 44; also Ann. Inst. Past., 43, 1929, 1658; B. ontarioni Paillot, B. presumably indicates Bacterium, see inde.K, p. 522, L'infection chez les insectes, 1933, 134; Steinhaus, Bacteria Associated Extra- cellularly with Insects, Minneapolis, 1942, 72.) From diseased larvae of the corn borer {Pyraustra nubilalis Hb.). Bacillus oogenes Migula. (Bacillus oogenes hydrosulfureus a, Zorkendorfer, Arch, f . Hyg., 16, 1893, 385 ; Migula, Syst. d. Bakt., 2, 1900, 573.) From hens' eggs. Bacillus opacus Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 214.) From bean infusions and fermenting cabbage. Bacillus orae Werner. (Cent, f . Bakt., II Abt., 87, 1933, 464.) Weak growth on agar containing calcium salts of formic, acetic, and butyric acids. One culture isolated from European soil. Bacillus oxylaclicus Dyar. (Dyar, Ann. N. Y. Acad. Sci., 8, 1895, 369; Bac- terium oxylacticus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 107.) From air and a culture from Krai's laboratory labeled Bacillus oxylacticus. The latter is given in the Krai 1900 catalogue as Bacillus oxalaticus Zopf and undoubtedly was the organism received by Migula from Zopf and studied by him (Migula, Arb. tech. Hochschule Karlsruhe, 1, Heft 1, 1894, 139 and Migula, Syst. d. Bakt., 2, 1900, 538). This is now regarded as having been Bacillus megatherium De Bary. Bacillus pabuli Schieblich. (Cent. f. Bakt., II Abt., 58, 1923, 204.) Com- monly isolated from green and ferment- ing fodder. Bacillus pallidus Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 98.) One strain isolated from soil from New York State. Bacillus palustris Sickles and Shaw. (Jour. Bact., 28, 1934, 418; Rhodobacillus palustris Sickles and Shaw, Jour. Bact., 38, 1939, 241.) Decomposes the specific carbohydrate of pneumococcus type III. From swamp and other uncultivated soils. Bacillus palustris var. gelaticus Sickles and Shaw {loc. cit., 419). A variety that decomposes agar slightly. Found only once. Bacillus paucicutis Burchard. (Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1902, 27.) From rye bread. Bacillus pectocutis Burchard. (Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1902, 24.) From the air. Bacillus pelagicus Russell. (Bot. Gaz., 18, 1S93, 383.) From sea water and marine mud from Woods Hole, Massa- chusetts. Bacillus pellucidus Soriano. (Revista FAMILY BACILLACEAE 751 del Inst it. Bacteriol., Buenos Aires, 6, 1935, 567.) Author saj's colonies resem- ble Bacillus simplex. Habitat probably in soil. Bacillus peptogenes (Buchanan and Hammer) Bergey et al. {Bacterium peptogenes Buchanan and Hammer, Iowa Agr. Exp. Sta. Res. Bull. 22, 1915, 273; Bergey et al., Manual, 1st ed., 1923, 293.) From a tube of litmus milk after auto- claving. Bacillus pcptonans Chester. {Bacil- lus lactis peptonans Sterling, Cent. f. Bakt., II Abt., 1, 1895, 473; Chester, Man. Determ Bact., 1901, 271.) From milk. Very similar to Bacillus mesen- tericus vulgatus Flligge. Bacillus peptonificans Lubenau. (Cent. f. Bakt., I Abt., Orig., 40, 1906, 435.) Similar to Bacillus subtilis. Be- lieved to be the cause of an epidemic of gastroenteritis. Bacillus perlucidulus Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 43.) Isolated 3 times from garden air. Bacillus petiolaius Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 48.) Isolated twice from garden air. Bacillus phaseoli von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 500.) From canned beans. Bacillus phenolphilos Bartels. (Cent, f. Bakt., II Abt., 103, 1940, 21.) Good growth on media containing m/50 phenol . One culture isolated from soil. Bacillus picrogenes Patrick and Werk- man. (Iowa State Coll. Jour. Sci., 7, 1933, 410.) Ferments xylan. One cul- ture isolated from decayed watermelon. Bacillus pilijormis Tyzzer. (Jour. Med. Research, 37, 1917, 307.) All at- tempts to cultivate the organism failed except when mixed with a streptotoccus. Considered aerobic by Ford (Textbook of Bact., 1927, 712). Causes fatal disease of Japanese waltzing mice. Bacillus piscicidus Migula. {Bacillus piscicidus agilis Siebert, Gazetalekarska, 1895, No. 13-17; abst. in Cent. f. Bakt., 17, 1895, 888; Bacierium piscicidus agilis Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 140; Migula, Syst. d. Bakt., 2, 1900, 652; Bacillus piscicidus nobilis (sic) Babes and Riegler, Cent, f. Bakt., II Abt., Orig., 33, 1902-03, 440.) Cause of a fish epidemic in St. Petersburg. Bacillus pisi von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 502.) From young peas. Bacillus plahjchoma Gray and Thorn- ton. (Cent. f. Bakt., II Abt., 73, 1928, 93.) Phenol is attacked. Three strains isolated from soil. Bacillus plexiformis Goadby. (Dental Cosmos, 42, 1900, 322.) From the mouth. Bacillus plicatus Deetjen. (Deetjen, Inaug. Diss., Wlirzburg, 1890; not Bacil- lus plicatus Frankland and Frankland, Philos. Trans. Royal Soc. London, 178, B, 1887, 273; not Bacillus plicatus Zim- mermann, Bakt. unserer Trink- u. Nutz- wasser, Chemnitz, I Reihe, 1890, 54; not Bacillus plicatus Copeland, Rept. Filtra- tion Commission, Pittsburgh, 1899, 348; not Bacillus plicatus Chester, Man. Determ. Bact., 1901, 275.) From sausage. Bacillus pollacii Pavarino. (Atti R. Accad. Naz. Lincei Rend. CI. Sci. Fis. Math, e Nat., 20, 1911, 233.) Reported to cause depressed spots on leaves of Odontoglossum citrosmum. Bacillus popxdi Brisi. (Atti Cong. Nat. Ital. Pom. della Soc. Ital. di Sci. Nat. Milano, 1907, 376.) Reported as cause of galls on branches of poplar trees {Populus sp.). Bacillus pseudanthracis Kruse. (Milz- brandahnlicher Bacillus, Burri, Hyg. Rundschau, 4, 1894, 339; abst. in Cent. f. Bakt., 16, 1894, 374; Kruse, in Flugge, Die Mikroorganismen, 3 Aufl., 2, 1896, 233; not Bacillus pseudanthracis Wahr- lich, Bakteriol. Studien, Petersburg, 1890-91, 26; Bacillus pseudo-anthracis Chester, Man. Determ. Bact., 1901, 280.) From South American bran. Bacillus pseudococcus Migula. (Bacil- 752 MANUAL OF DETERMINATIVE BACTERIOLOGY lus No. 11, Pansini, Arch. f. path. Anat., 122, 1890, 446; Migula, Syst. d. Bakt., 2, 1900, 557.) From sputum. Bacillus pseudodiphthe7'icum magnus Pdegaard. (Acta Path, et Microbiol. Scand., 21, 1944, 451; see Endicott, Biol. Abst., 20, 1946, 12926.) From the nose of a child suspected of having diphtheria. Resembles Corynehacterium diphthcriae in young cultures. Non-pathogenic. Bacillus pseudofusiformis Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 47.) Isolated once from garden air. Bacillus pseudosubtilis Migula. (Bacillus subtil is similis Sternberg, Manual of Bact., 1893, 679; Migula, Syst. d. Bakt., 2, 1900, 618.) From the liver of a yellow fever cadaver. Bacillus punctiformis Chester. (Bacil- lus No. 23, Conn, Storrs Agr. Exp. Sta. Rept., 1893, 53; Chester, Man. Determ. Bact., 1901, 284.) From milk. Bacillus pycnoticus Grohmann . (Cent . f. Bakt., II Abt., 61, 1924, 261 ; Ruhland and Grohmann, Jahrb. wissensch. Botanik, 63, 1924, 321 ; Hydrogenomonas pycnotica Bergey et al.. Manual, 3rd ed., 1930, 34.) Oxidizes hydrogen in the presence of oxygen to form water. Pre- sumably widely distributed in soil. Bacillus quercifolius Deetjen. (Deet- jen, Inaug. Diss., Wiirzburg, 1890; Bacterium quercifolium Migula, Syst. d. Bakt., 2, 1900, 309.) From sausage. Bacillus rarerepertus Schieblich. (Cent. f. Bakt., I Abt., Orig., 124, 1932, 269.) From beet leaves. Bacillus rarus Werner. (Cent. f. Bakt., II Abt., 87, 1933, 456.) Good growth on Ca n-butyrate agar. One culture isolated from forest soil of Germany. Bacillus repens Gibson. (Cent. f. Bakt., II Abt., 92, 1935, 370.) Decom- poses urea. Eight strains isolated from soil. Bacillus reptans Ghosh. (Compt. rend. Soc. Biol., Paris, 86, 1922, 914.) From a case of appendicitis. Bacillus retaneus Grohmann. (Mor- phologisch- biologische Beitrage zur Kenntnis der Wasserstoffbakterien, In- aug. Diss., Univ. Leipzig, 1923; Cent. f. Bakt., II Abt., 61, 1924, 267 ; Ruhland and Grohmann, Jahrb. wissensch. Botanik, 63, 1924, 321.) Oxidizes hydrogen in the presence of oxygen to form water. Pre- sumably widely distributed in soil. Bacillus retijormis Migula. (Netz- bacillus, Maschek, Bakt. Untersuch. d. Leitmeritzer Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 712.) From water. Bacillus robust us Weiss. (Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 247; not Bacillus robuslus Blau, Cent. f. Bakt., II Abt., 15, 1905, 134.) From fermenting beets. Bacillus ruber Zimmermann. (Zim- mermann. Die Bakt. unserer Trink- u. Nutzwasser, Chemnitz, 1, 1890, 24; not Bacillus ruber Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 3, 1875, 181; Bacillus pseudoruber Migula, Syst. d. Bakt., 2, 1900, 850; Eryihrobacillus ruber Holland, Jour. Bact., 5, 1920, 223, line 15; Serratia rubra Bergey et al.. Manual, 1st ed., 1923, 92; Chromobacterium ruber Topley and Wilson, Princip. of Bact. and Im- mun., 1, 1931, 402.) From Chemnitz tap water. Spherical spores. See Manual , 5th ed . , 1 939, 607 for a description of this species. Bacillus rufescens Sttihrk. (Cent. f. Bakt., II Abt., 93, 1935, 178.) Charac- terized by good growth on Ca n-butyrate agar. One culture isolated from garden soil of Germany. Bacillus rufulus Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 59.) Isolated 3 times from garden air. Bacillus rugosus Henrici. (Henrici, Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 28; not Bacillus rugosus Wright, Mem. Nat. Acad. Sci., 7, 1895, 438; not Bacillus rugosus Chester, Man. Determ. Bact., 1901, 220.) From Swiss cheese. Bacillus rugulosus Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 181.) One of a FAMILY BACILLACEAE 753 group of species described as growing well on Ca n-butyrate agar. Three strains isolated from soils of Germany, Cuba, and Italy. Bacillus sacchari Janse. (Mededeel. uit's Lands. Plantentuin, 9, 1891, 1.) Reported to cause sereh, a disease affect- ing sugar cane {Saccharum officinarum). Went (Arch, voor de Java Suikerindus- trie, 1895, 589) regards this as probably Bacillus subtilis. Bacillus saccharolyticus Nepomnjat- schjaja and Libermann. (Jour. f. Mikrobiol., Ukraine, S, 1938, 57; abst. in Cent. f. Bakt., II Abt., 101, 1940, 81.) From plum preserves. A gas-producing rod. Bacillus saccohranchi Dobell. (Quart. Jour. Micro. Sci., 56, 1911, 441.) From heart blood of a fish {Saccobrnrichus fossilis) . Bacillus santiagensis Stiihrk. (Cent. f. Bakt., II Abt., 9S, 1935, 188.) Good growth on Ca n-butyrate agar. One culture isolated from Cuban soil. Bacillus saprogenes Migula. (Bacillus saprogenes vini III, Kramer, Bakteriolo- gie in ihren Beziehungen zur Landwirt- schaft, 2, 1892, 137; Migula, Syst. d. Bakt., 2, 1900, 572; not Bacillus sapro- genes I, II and ///, Herfeldt, Cent. f. Bakt., II Abt., 1, 1895, 77; not Bacillus saprogenes Salus, Arch. f. Hyg., 51, 1904, 115.) From wine. Bacillus saprogenes Chester. (Bacil- lus saprogenes vini VI, Kramer, Bak- teriol. Landwirtsch., 2, 1892, 139; Ches- ter, Man. Determ. Bact., 1901, 289; not Bacillus saprogenes Trevisan, I generi e le specie delle Batteriacee, 1889, 17.) From wine. Bacillus scaber (Duclaux) Trevisan. (Tyrothrix scaber Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From milk. Bacillus schottelii Trevisan. (Darm- bacillus, Lydtin and Schottelius, Der Rotlauf der Schweine, Weisbaden, 1885, 214; Bacillus coprogenes foetidus Flligge, Die Mikroorganismen, 2 Aufl., 1886, 305; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 17; Bacterium coprogenes Migula, Syst. d. Bakt., 2, 1900, 327; Bacterium schottelii Chester, Man. De- term. Bact., 1901, 197.) From the intestinal contents of swine. Bacillus segetalis Werner. (Cent. f. Bakt., II Abt., 87, 1933, 467.) Charac- terized by growth on Ca n-butyrate agar. One strain isolated from soil in Germany. Bacillus septico-aerobius Hauduroy et al. (Bacille septique a^robie, Legros, These M(5d., Paris, 1902 ; Hauduroy et al., Diet. d. Bact. Path., 1937, 46.) Aerobic, facultative. From a case of acute gaseous gangrene. Bacillus seplicus insectorum Krassil- stschik. (Memoires de la Soc. Zool. de France, 6, 1893, 250.) From cockchafer larvae (Melolontha melolontha). Bacillus serrulatus Stiihrk. (Cent. f. Bakt., II Abt., 9S, 1935, 193.) Only moderate growth on Ca n-butyrate agar. One culture isolated from Cuban soil. Bacillus sesami Malkoff. (Cent. f. Bakt., II Abt., 16, 1906, 665.) Patho- genic on sesame (Sesamum). Bacillus siccus Chester. (Bacillus No. 25, Conn, Storrs Agr. Exp. Sta. Kept., 1893, 63; Chester, Man. Determ. Bact., 1901,284.) From milk. Bacillus similis Schroeter. (Bacillus II, Bienstock, Ztschr. f. klin. Med., 8, 1884, Heft 1 and 2; Schroeter, in Cohn, Kryptog. -Flora v. Schlesien, 3, 1, 1886, 160; Bacillus coprocinus Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 15; Bacillus faecalis No. II, Kruse, in Flligge, Die Mikroorganismen, 3 Aufl., 2, 1896, 215; Bacterium simile Chester, Man. Determ. Bact., 1901, 197.) From feces. Bacillus similis Eckstein. (Arch. f. Forst- u. Jagdwesen, 26, 1894, 11.) From infected larvae of the nun moth (Lyman- tria monacha), etc. Bacillus similityphosus Migula. (Ty- phusahnlicher Bacillus, Maschek, Bakt. Untersuch. d. Leitmeritz. Trinkwasser, Leitmeritz, 1887; Migula, Syst. d. Bakt., 2, 1900, 730. ) From water. 754 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacillus sinapivagus Kossowicz. (Cent. f. Bakt., II Abt., 28, 1909, 241.) From pickles. Bacillus sombrosus Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 429.) From the stomach of a bird. Bacillus sorghi Burrill. (The Micro- scope, 7, 1887, 321; Proc. Amer. Soc. Microscopists, 1888, 193; Bacterium sorghi Chester, Delaware Agr. E.xp. Sta. Ann. Rept., 9, 1897, 127; Elliott and Smith, Jour. Agr. Res., 38, 1928, 1.) Reported to cause a disease of sorghum {Holcus sorghum). Bacillus sotlo Paillot. (Sotto-Bacillus, Ischivata, quoted from Aoki and Chi- gasaki, Mitteil. d. Med. Fakul. d. k. Univ. z. Tokyo, 13, 1915, 419 and U, 1915, 59; Bacterium sotto Metalnikov and Chorine, Internat. Corn Borer Invest., Sci. Repts., 1, 1928, 56; Paillot, ibid., 1, 1928, 77-106.) From silkworms {Bombyx viori). Sotto is Japanese for "plotzlich hinf alien". Bacillus spatiosus Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 56.) Isolated once from garden air. Bacillus spermatozoides Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 13.) From dead moths {Hyponomeuta evonym,ella) . Bacillus sphaerosporus Beijerinck. (Cent. f. Bakt., II Abt., 25, 1909, 45.) This organism has round terminal spores and produces nitrous oxide from nitrates. From garden soil. Bacillus sphaerosporus calco-aceticus Beijerinck {loc. cit., 46). A variety of the above having spherical to ellipsoidal spores. Bacillus spinosporus Soriano. (The- sis, Univ. Buenos Aires, 1935, 562.) Spores spinate like some strains of Bacillus pohjmyxa. No gas formed. One strain isolated from soil. Bacillus spiralis Migula. (Syst. d. Bakt., 2, 1900, 624.) From water. Bacillus spirogyra Dobell. (Quart. Jour. Micro. Sci., 53, 1909, 579 and 56, 1911, 434.) From large intestine of frog {Rana leniporaria) and toad {Bufo vul- garis) . Bacillus spongiosxis Aderhold and Ruh- land. (Cent. f. Bakt., II Abt., 15, 1905, 376.) Found in the gum masses dis- charged by cherry trees. Bacillus sporonemo.Schaudinn. (Arch. f. Protistenkunde, 2, 1903, 421.) From sea water. Bacillus spurius Grohmann. (Mor- phologisch-biologische Beitrage zur Kenntnis der Wasserstoffbakterien, In- aug. Diss., Univ. Leipzig, 1923; Cent. f. Bakt., II Abt., 61, 1924, 267; Ruhland and Grohmann, Jahrb. wissensch. Botanik, 63, 1924, 321.) Oxidizes hydro- gen in the presence of oxygen to form water. Presumably widely distributed in soil. Bacillus sputi Chester. (Bacillus No. 6, Pansini, Arch. f. path. Anat., 122, 1890, 442; Chester, Man. Determ. Bact., 1901,280.) From sputum. Bacillus squajniformis Saito. (Jour. Coll Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 54.) Isolated 9 times from garden soil. Bacillus stellaris Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 52.) Isolated 6 times from garden air. Bacillus stellatus Zimmermann. (Zim- mermann, Bakt. unserer Trink- u. Nutz- wasser, II Reihe, 1894, 14; not Bacillus stellatus Chester, Man. Determ. Bact., 1901, 274; not Bacillus stellatus Vincent, Ann. Inst. Past., 21, 1907, 62.) From water. Bacillus sireptoformis Migula. (Sal- peter zerstorenden Bacillus, Schirokikh, Cent. f. Bakt., II Abt., 2, 1896, 204; Migula, Syst. d. Bakt., 2, 1900, 682; Bacillus schirokikhi Chester, Man. De- term. Bact., 1901, 252; Bacillus denitri- ficans Chester, ibid., 274.) From horse feces. Utilizes potassium nitrate. Bacillus suaveolens Sclavo and Gosio. (Quoted from Omeliansky, Jour. Bact., 8, 1923, 398.) No source given. Bacillus subcuticularis Migula. (Bacillus cuticularis albus Tataroff, In- aug. Diss., Dorpat, 1891, 24; Migula, Syst. d. Bakt., 2, 1900, 623; Bacillus cuticularis Chester, Man. Determ. Bact.^ ie01,285.) From water. FAMILY BACILLACEAE 755 Bacillus sublanatus Wright. (Mem. Nat. Acad. Sci., 7, 1895, 455.) From water. Bacillus sublustris Schieblich. (Cent, f. Bakt., II Abt., 58, 1923, 206.) Com- monly isolated from green and fermenting fodders . Bacillus submarinas ZoBell and Up- ham. (Bull. Scripps Inst, of Oceanogra- phy, Univ. Calif., 5, 1944, 267.) Central ovate spores. From marine bottom deposits. Bacillus (Streptobacter) subtiliformis Schroeter. (Bacillus I, Bienstock, Ztschr. f. klin. Med., 8, Heft 1 and 2, 1884; Schroeter, in Cohn, Kryptog.- Flora V. Schlesien, 3, 1, 1886, 160; Bacil- lus mesenthericus (sic) Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 15; not Bacillus mesenlericus Trevisan, ibid., 19; Bacillus subtilis sim7tlans I, Eisenberg, Bakt. Diag., 3 Aufl., 1891, 189; Bacillus faecalis No. I, Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., f, 1896, 215; Bacterium subtile Migula, Syst. d. Bakt., 3, 1900, 292; Bacterium subtiliforme Chester, Man. Determ. Bact., 1901, 197.) From feces. Bacillus succinicus Fitz. (Quoted from DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 966.) From infusions. Bacillus suffuscus Grohmann. (i\Ior- phologisch-biologische Beitrage zur Kenntnis der Wasserstoffbakterien, In- aug. Diss., Univ. Leipzig, 1923; Cent. f. Bakt., II Abt., 61, 1924, 267;RuhIand and Grohmann, Jahrb. wissensch. Botanik, 63, 1924, 321.) Oxidizes hydro- gen in the presence of oxygen to form water. Presumably widely distributed in soil. Bacillus supraresistans Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 185.) Very good growth on Ca n-butyrate agar. One culture isolated from soil in Germany. Bacillus surgeri Dornic and Daire. (Bull, mensuel de 1 'Office de renseigne- ments agricoles, 6, 1907; abst. in Rev. Gen. du Lait., 6, 1907, 164.) Spores not observed but author stated that they were probably present because this species could withstand 85°C for 5 minutes. From whey. Bacillus tabaci III, Koning. (Tijdschr. voor toegepaste scheikunde en hygiene. Deel 1, 1897. See Behrens, Mykologie der Tabakfabrikation, in Lafar, Hand- buch der techn. Mykologie, 5, 1905, 11.) Thermophilic. Probably from soil. Bacillus tardivus Stiihrk. (Cent. f. Bakt., II Abt., 93, 1935, 177.) Very slight growth on Ca n-butyrate agar. One culture isolated from garden soil of Germany. Bacillus technicus Morikawa and Pres- cott. (Jour. Bad., 13, 1927, 58; also see Morikawa, Bull. Agr. Chem. Soc. Japan, 3, 1927, 28.) Produces butyl and iso- propyl alcohols. Source not given. Bacillus tenax Eckstein. (Ztschr. f. Forst- u. Jagdwesen, 26, 1894, 14.) From larvae of the nun moth {Lymantri a monacha). Bacillus tenuis non-liquefaciens Chou- kevitch. (Ann. Inst. Past., :?5, 1911, 352.) From large intestine of horse. Bacillus ierminalis Migula. (Bacillus No. XII, Flugge, Ztschr. f. Hyg., 17, 1894, 296 ; Bacillus lactis No. XII, Kruse, in Flugge, Die Mikroorganismen, 3 Aufl., 2, 1896, 269; Migula, Syst. d. Bakt., 2, 1900, 578; Bacillus lacteus Chester, Man. Determ. Bact., 1901, 291.) From milk. A duplicate of Bacterium semper- vivum Migula. Bacillus terminalis var. thermophilus Prickett. (N. Y. Agr. Exp. Sta. Tech. Bull. 147, 1928, 44.) Produces a brown water soluble pigment on agar ; optimum temperature 55°C to 65°C. Fourteen strains from raw and pasteurized milk, milk powder, and skin of a cow. Bacillus terrestris Werner. (Cent. f. Bakt., II Abt., 87, 1933, 461.) Weak growth on Ca n-butyrate agar. Two strains isolated from soils of Germany. Bacillus ietanoides Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 49.) Isolated once from garden air. Bacillus thalassokoites ZoBell and Up- ham. (Bull. Scripps Inst, of Oceanogra- phy, Univ. Calif., 5, 1944, 268.) Central spores. From marine bottom deposits. 756 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacillus theae Hori and Bokura. (Jour. Plant Protection, Tokyo, 2, 1915, 1.) Pathogenic for tea {Thea sinensis). Bacillus thermoahundans Beaver. (Dissertation, Ohio State University, Columbus, 1932, 31.) Thermophilic, sub- terminal spores. Growth at 55°C, less growth at 37°C. From malted milk powder. Bacillus thermoacetigenitus Beaver, loc. cit., 25. Thermophilic, central spores. No growth at 37°C. From vinegar. Bacillus thermoacidificans Renco. (Ann. Microbiol., 2, 1942, 000.) From Grana cheese whey. This is stated by Gorini (R. 1st. Lombardo Sci. e. Lett., Rend., 76, 7« della Ser. 3, 1942, 3) to be the same as Bacillus laciis termophilus Gorini. Bacillus thermoactivus Beaver, loc. cit., 27. Thermophilic, central spores. No growth at 37°C. From home-canned beets. Bacillus thermoannulatus Beaver, loc. cit., 17. Thermophilic, subterminal spores. No growth at 37°C. From com- mercially canned tomatoes. Bacillus thermoaquatilis Beaver, loc. cit., 18. Thermophilic, subterminal spores. No growth at 37°C. From a warm spring at Springfield, Ohio. Bacillus thermoarborescens Beaver, loc. cit., 30. Thermophilic, sub-terminal to central spores. Growth at 55°C, less growth at 37°C. From candy. Bacillus thermobutyrosus Beaver, loc. cit., 15. Thermophilic, subterminal spores. No growth at 37°C. From com- mercially canned tomatoes. Bacillus thermocompactus Beaver, loc. ict., 20. Thermophilic, subterminal spores. No growth at 37°C. From red grapes stored in sawdust. Bacillus thermodaclylogenitus Beaver, loc. cit., 28. Thermophilic, central to subterminal spores. Growth at 37°C and 55°C. From commercially packed dates. Bacillus thennoejfervescens Beaver, loc. cit., 23. Thermophilic, central spores. No growth at 37°C. From commercially canned peas. Bacillus thermofaecalis Beaver, loc. cit., 30. Thermophilic, subterminal spores. Growth at 55°C. From feces of robin. Bacillus thermojibrincolus Itano and Arakawa. (Ber. d. Ohara Inst. f. land- wirsch. Forschungen, Japan, 4, 1929,265.) Thermophilic; decomposes cellulose. From soil containing decomposed leaves. Bacillus thermofiliformis Beaver, loc. cit., 22. Thermophilic, subterminal spores. No growth at 37°C. From com- mercially canned peas. Bacillus thermograni Beaver, loc. cit., 16. Thermophilic, subterminal spores. No growth at 37°C. From commercially canned corn. Bacillus thermolongus Beaver, loc. cit., 19. Thermophilic, subterminal spores. No growth at 37°C. From commercially canned tomatoes. Bacillus thermolubr leans Beaver, loc. cit., 26. Thermophilic, central spores. No growth at 37°C. From lubricating oil . Bacillus thermononodorus Beaver, loc. cit., 26. Thermophilic, central spores. No growth at 37°C. From tap water. Bacillus thermonubilosus Beaver, loc. cit., 19. Thermophilic, subterminal spores. No growth at 37°C. From soil. Yellow Springs, Ohio. Bacillus thennoodoratus Beaver, loc. cit., 29. Thermophilic, central spores. Growth at 55°, less growth at 37°C. From spoiled gelatin. Bacillus thermopellitus Beaver, loc. cit., 22. Thermophilic, central spores. No growth at 37°C. From old sour milk. Bacillus thermophilus Miquel. (Mi- quel, Ann. d. Microgr., 1, 1888, 4; Ba- cillus thermophilus miquelii Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 269; Bacterium termophilum (sic) Migula, Syst. d. Bakt., 2, 1900, 342; Bacterium miquelii Chester , Man. De- term. Bact., 1901, 186.) From the FAMILY BACILLACEAE iOt intestine, water and soil. Ther- mophilic. No growth below 40°C. Bacillus ihermophilus sojae Rokusho and Fukutome. (Jour. Agr. Chem. Soc, Japan, 13, 1937, 1235.) From spon- taneously heating soj'-bean cake. Bacillus thermosuavis Beaver, loc. cit., 24. Thermophilic, central spores. No growth at 37°C. From commercially- canned mincemeat. Bacillus thermotenax Beaver, loc. cit., 28. Thermophilic, subterminal spores. Growth at 37°C and 55°C. From ground horseradish. Bacillus thermourinalis Beaver, loc. cit., 16. Thermophilic, subterminal spores. No growth at 37°C. From hu- man urine. Bacillus thermouiscidus Beaver, loc. cit., 21. Thermophilic, subterminal spores. No growth at 37°C. From fresh pig ovary. Bacillus thoracis Howard. (Gleanings in Bee Culture, 28, 1900, 124.) From black brood of the honey bee (Apis mellifera). Bacillus tracheitis sive graphitosis Krassilstschik. (Memoires de la Soc. Zool. de France, 6, 1893, 2.50.) From diseased larvae of the cockchafer (Melo- lontha melolontha ) . Bacillus tricomii Trevisaa. (Bacillo della gangraena senilis, Tricomi, Riv. internaz. di Med. e Chir., 3, 1886, 73; Trevisan, I generi e le specie delle Bat- teriacee, 1889, 13; Bacterium tricomii Migula, Syst. d. Bakt., 2, 1900, 310.) From a case of senile gangrene. Bacillus trifolii Voglino. (Ann. R. Accad. Agr. Torino, 39, 1896, 85. ) Patho- genic for clover (Trifolium pratense, T . repens, T. resupinatum) . Bacillus tritus Batchelor. (Jour. Bact., 4, 1919, 29.) One culture from feces. Bacillus tuberis von Wahl. (Cent. f. Bakt., II Abt., 16, 1906, 503.) From cooked truffles {Tuber oestivum). Bacillus tuberostis Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 248.) From fermenting beets. Bacillus tubifex Dale. (Annals of Bot., 26, 1912, 133.) Reported to cause a leaf disease of potato {Solanum tuberosum) and tomato {Lycopersicum esculentum). Bacillus turgidus (Duclaux) Trevisan. {Tyrothrix turgidus Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23; Trevisan, I generi e le specie delle Batteriacee, 1889, 16.) From milk. Bacillus tympani-cuniculi Morcos. (Jour. Bact., 23, 1932, 454.) Causes tympanitis in young rabbits. Bacillus ubicuitarius Soriano. (Es- tudio sistematico de algunas bacterias esporuladas aerobias. Thesis, Univ. Buenos Aires, 1935, 569.) Four cultures isolated from soil. Bacillus ulna Cohn. (Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 2, 1872, 177; also see Prazmowski, Untersuch. ii. d. Ent- wickelungsges. u. Fermentwirk. einiger Bakterienarten, Leipzig, 1880, 20.) Found once in an infusion of cooked egg-white. Bacillus undulatus den Dooren de Jong. (Bull. Assoc. Diplomes de Micro- biol. Nancy, No. 26-27, 1946, 12.) From soil. Bacillus uvaeformis Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 415.) From the stomachs and intestines of birds. Bacillus vaculatus Ravenel. (Mem. Nat. Acad. Sci., 8, 1896, 31.) From soil. Bacillus validus Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 97.) Four cultures isolated from soil from Ger- many, Cuba, and Egypt. Bacillus valinovorans Heigener. (Cent. f. Bakt., II Abt., 93, 1935, 104.) Good growth on valine agar. Five strains isolated from soils from Egypt, Germany, Italy, and Palestine. Bacillus varians Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 50.) Isolated 11 times from gar- den air. Bacillus ventricosus Heigener. (Cent. 758 MANUAL OF DETERMINATIVE BACTERIOLOGY f. Bakt., II Abt., 93, 1935, 102; not Bacillus ventricosus Weiss, Arb. bakt. Inst. Karlsruhe, 2, 1898, 233.) One culture isolated from soil from Italy. Bacillus ventriculus Koch. (Botan. Zeitung, 46, 1888, 341.) From slices of carrot exposed to the air. Formed two spores in a spindle-shaped sporangium. Bacillvs vernicosus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 46; not Bacil- lus vernicosus Migula, Syst. d. Bakt., 2, 1900, 781.) From waste water. Bacillus verticillatus Ravenel. (Rav- enel, Mem. Nat. Acad. Sci., 8, 1896, 13; Bacterium verticillatum Chester, Man. Determ. Bact., 1901, 192.) From soil. Bacillus vesicae Migula. {Bacillus sepiicus vesicae Clado, quoted from Eisenberg, Bakt. Diag., 3 Aufi., 1891, 341 ; Migula, Syst. d. Bakt., 2, 1900, 620.) From urine in a case of cystitis. Prob- ably is Bacillus cladoi Trevisan. Bacillus villosus Zimmermann. (Bakt. unserer Trink- u. Nutzwasser, Chemnitz, II Reihe, 1894, 38; not Bacillus villosus Keck, Inaug. Diss., Dorpat, 1890, 47.) From water. Bacillus violaceus Eisenberg. (Bakt. Diag., 2 Aufl., 1888, 8; not Bacillus vio- laceus Schroeter, Kryptogamen-Flora von Schlesien, 3, 1886, 157; not Bacillus violaceus Frankland and Frankland, Ztschr. f. Hyg., 6, 1888, 394.) Said to produce central spores. From water. Bacillus viridi-glaucescens Sack. (Cent. f. Bakt., II Abt., 65, 1925, 113.) From several kinds of soil . Bacillus viridiluteus Pagliani et al. (Griingelber, nicht verfliissiger Bacillus, Eisenberg, Bakt. Diag., 1 Aufl., 1886, Tab. 6; Pagliani, Maggiora and Fratini, Soc. ital. d'igiene, 1887,586, see Trevisan, I generi e le specie delle Batteriacee, 1889, 19). From water. Bacillus viscosus hruxellensis van Laer. (Cent. f. Bakt., II Abt., 23, 1909, 159.) From beer wort. Bacillus viticola Burgwitz. (Bacillus vitis Merjanian and Kovaleva, Prog. Agric. et Vitic, 95, 1930, 594 and 96, 1931, 17; not Bacillus vitis Montemartiui, Rev. Patol. Veg., 6, 1913, 175; Burgwitz, Phytopath. Bacteria, Leningrad, 1935, 37.) Pathogenic for the grape vine. Bacillus vitreus Migula. (No. 11, Lembke, Arch. f. Hyg., 26, 1896, 306; Migula, Syst. d. Bakt., 2, 1900, 569.) From the intestines of infants. Bacillus vogelii Migula. (Roter Kart- off el bacillus, Vogel, Ztschr. f. Hyg., 26, 1897, 404; Migula, Syst. d. Bakt., 2, 1900, 556 ; Bacillus viscosus Chester, Man. Determ. Bact., 1901, 286; not Bacillus viscosus Frankland and Frankland, Ztschr. f. Hyg., 6, 1889, 391.) From stringy bread. Bacillus watzmannii Werner. (Cent, f . Bakt., II Abt., 87, 1933, 462.) Weak growth on Ca n-butyrate agar. One culture isolated from soil of Germany. Bacillus weigvianni Migula. (Bak- terie II, Weigmann and Zirn, Cent. f. Bakt., 15, 1894, 465; Migula, Syst. d. Bakt., 2, 1900, 693.) From soapy milk. Bacillus xylanicus Patrick and Werk- man. (Iowa State Coll. Jour. Sci., 7, 1933, 415.) Ferments xylan. One cul- ture isolated from decayed apple wood. Bacillus xylophagus Patrick and Werk- man. (Iowa State Coll. Jour. Sci., 7, 1933, 414.) Ferments xylan. One cul- ture isolated from decayed apple wood. Bacillus zirnii Migula. (Bakterie III, Weigmann and Zirn, Cent. f. Bakt., 15, 1894, 466; Migula, Syst. d. Bakt., 2, 1900, 693.) From soapy milk. Bacterium adametzii Migula. (Bacil- lus No. XIV, Adametz, Landwirtsch. Jahrb., 18, 1889, 246; Migula, Syst. d. Bakt., 2, 1900, 338; Bacterium rugosum Chester, Man. Determ. Bact., 1901, 194.) From cheese. Bacterium aloes Passalacqua. (Rev. Pat. Veg., 19, 1929, 110.) From diseased aloes. Bacterium angulans Burchard. (In- aug. Diss., 1897; Arb. bakt. Inst. Karls- ruhe, 2, Heft 1, 1902, 43.) From water. Bacterium aqueum Migula. (Bacillus thertnophilus VIII, Rabinowitsch, Ztschr. f. Hyg., 20, 1895, 160; Migula, FAMILY BACILLACEAE 759 Syst. d. Bakt., 2, 1900, 345; Bacterium thermophilum VIII, Chester, Man. Determ. Bact., 1901, 186.) From feces and corn. Bacterium articulatum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1897, 445.) From the stomach and intestines of birds. Bacterium asparagi von Wahl. (Von Wahl, Cent. f. Bakt., II Abt., 16, 1906, 498; Bacillus asparagi Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 436.) From boiled asparagus. Bacterium, brachysporum Burchard. (Inaug. Diss., 1897; Arb. bakt. Inst. Karlsruhe, 2, Heft 1, 1902, 20.) From bakery bread. Bacterium canadensis Chorine. (In- ternat. Corn Borer Invest., Sci. Rpts., 2, 1929, 39; also Ann. Inst. Past., 43, 1929, 1658; Bac. canadensis Chorine and Metal - nikov, Ann. Inst. Past., 43, 1929, 1392; also Paillot, L'infection chez les insectes, 1933, 134 where Bac. equals Bacterium, see index p. 522; Bacillus canadensis Steinhaus, Bacteria Associated Hxtra- cellularly with Insects, Minneapolis, 1942, 50.) In its general characters said to resemble Bacillus megatherium and other bacteria isolated from insects {Bacillus thnringiensis, Bacillus hoplo- sternus, etc.). Pathogenic for larvae of Pyrausta nuhilalis, Galleria mellonella, and Epheslia kuhniella. From disea.sed larvae of the corn borer. Bacterium caitleyae Pavarino. (Atti R. Accad. Xaz. Lincei Rend. CI. Sci. Fis., Mat. e. Nat., 20, 1911, 233.) From diseased orchids. Bacterium cazauhon Metalnikov. (Compt. rend. Soc. Biol., Paris, 105, 1930, 536; two varieties, Bacterium cazauhon I and II, are recognized by Metalnikov, Ermolaev and Schobaltzyn, Internat. Corn Borer Invest., Sci. Repts., 3, 1930, 30 and Ann. Inst. Past., 46, 1931, 469.) From diseased corn borer larvae {Pyrausta nuhilalis Hb.). Bacterium christiei Chorine. (Inter- nat. Corn Borer Invest., Sci. Rpts., 2, 1929, 46; also Ann. Inst. Past., 43, 1929, 1666.) According to the author, this closely resembles Bacterium ontarioni. Several strains isolated from diseased corn borers. Bacterium colomatii Chester. (Colo- matii, Breslauer arztliche Ztschr., 1883, No. 4; Chester, Man. Determ. Bact., 1901, 186.) From xerotic masses in conjunctivitis. Bacterium deliense Swellengrebel. (Archiv f. Protist., 31, 1913, 277.) Ob- served in stained smears from the spleen of diseased cattle but not isolated. Two spores may form in a single cell if division is delayed. Bacterium ephestiae No. 1 and No. 2 ^letalnikov and Chorine. (Ann. Inst. Past., 43, 1929, 1394.) Not pathogenic for corn borer although the size of the larvae was reduced. Later, Ellinger and Chorine (Internat. Corn Borer Investigations, Sci. Rpts., 3, 1930, 37) identified these as strains of Bacillus thuringiensis. From diseased larvae of Epheslia kuehniella. Bacterium filiforme Henrici. (Hen- rici, Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 41; not Bacterium filiforme Migula, Syst. d. Bakt., 2, 1900, 296; Bacterium suhfiliforme Migula, ihid., 297.) From Swiss cheese. Bacterium galleriae No. 1, Chorine. (Batonnet mince, Metalnikov, Compt. rend. Acad. Sci., 'Paris, 175, 1922, 69; Chorine, Ann. Inst. Past., 4i, 1927, 1115.) From diseased larvae of the bee moth {Galleria mellonella). Bacterium galleriae Chorine. (Plus grand batonnet, ^letalnikov, Compt. rend. Acad. Sci., Paris, 175, 1922, 70; Chorine, Compt. rend. Soc. Biol., Paris, 95, 1926, 200; Bacterium galleriae No. 2, Chorine, Ann. Past. Inst., 41, 1927, 1117.) From diseased larvae of the bee moth {Galleria mellonella) . Resembles Bacil- lus megatherium. Pathogenic for the corn borer (Internat. Corn Borer Invest., Sci. Repts., 1, 1927,46). Bacterium galleriae No. 3, Chorine. (Ann. Inst. Past., 41, 1927, 1118.) From diseased larvae of the bee moth {Galleriae mellonella). Resembles Bacillus subtilis and Bacillus mesentericus. 760 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacterium giganteum Kern. (Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1896, 453.) From the stomach and intestines of birds. Bacterium glaucescens Migula. {Ba- cillus thermophilus VI, Rabinowitsch, Ztschr. f. Hyg., £0, 1895, 158; Migula, Syst. d. Bakt., 2, 1900, 344; Bacterium thermophilum VI, Chester, Man. De- term. Bact., 1901, 185.) From feces. Bacterium glutinosum. Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 441.) From the stomach of a dove. Bacterium ilidzense Migula. {Bacillus ilidzensis capsulatus Karlinski, Hy- gienische Rundschau, 5, 1895, 688; Migula, Syst. d. Bakt., 2, 1900, 340.) From the water of a hot spring. Ther- mophilic. Bacterium implectans Burchard. (In- aug. Diss., 1897; Arb. bakt. Inst. Karls- ruhe, 2, Heft 1, 1898, 29.) From drink- ing water. Bacterium, insulosuin Wilhelmy. (Arb. bakt. Inst. Karlsruhe, S, 1903, 16.) From meat extract. Bacterium insulum Weiss. (Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 252.) From fermenting malt. Bacterium, intactum Migula. {Bacil- lus thermophilus V, Rabinowitsch, Ztschr. f. Hyg., 20, 1895, 158; Migula, Syst. d. Bakt., 2, 1900, 344; Bacterimn thermophilum V , Chester, Man. Determ. Bact., 1901, 185.) From feces and corn. Bacterium iris Migula. (Irisierender Bacillus, Tataroff, Inaug. Diss., Dorpat, 1891, 57; Migula, Syst. d. Bakt., 2, 1900, 313; not Bacterium iris Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 125.) From water. Bacterium italicuni No. 1 and No. 2, Metalnikov, Ermolaev and Skobaltzyn. (Ann. Inst. Past., 46, 1931, 470; No. 2 is also described in Internat. Corn Borer Invest. Sci. Repts., 3, 1930, 30.) From larvae of the corn borer {Pyrausta nubilalis ) . Bacterium longum Kern. (Kern, Arb. bakt. Inst. Karlsruhe, /, Heft 4, 1894, 391; Bacterium squamosum longum Kern, ibid., 458; Bacillus squamosus longus Chorine, Ann. Inst. Past., 41, 1927, 1114.) From the intestines of a dove {Columba oenas). Bacterium lunula Dobell. (Quart. Jour. Micro. Sci., 53, 1909, 579.) From rectum of the toad {Bufo vulgaris). Resembles Bacterium hinnclealum Swel- lengrebel. Bacterium lydiae Migula. {Bacillus thermophilus I, Rabinowitsch, Ztschr. f. Hyg., 20, 1895, 156; Migula, Syst. d. Bakt., 2, 1900, 343; Bacterium ther- mophilum I, Chester, Man. Determ. Bact., 1901, 185.) Widely distributed in soil, snow, feces, corn. Bacterium m,ansfieldii Chester. (Ba- cillus No. 18, Conn, Storrs Agr. Expt. Sta., 1893, 51; Chester, Man. Determ. Bact., 1901, 197.) From milk. Bacterium markusfeldii Chester. (Ba- cillus der trichorrhexis nodosa, Markus- feld. Cent. f. Bakt., I Abt., 21, 1897, 230; Chester, Man. Determ. Bact., 1901, 192. ) Associated with the disease, tricho- rrhexis nodosa. Bacterium mesentericumMigula. {Ba- cillus mesentericus panis viscosi I, Vogel, Ztschr. f. Hyg., 26, 1897, 404; Migula, Syst. d. Bakt., 2, 1900, 314; Bacterium panis Chester, Man. Determ. Bact., 1901, 196.) From stringy bread dough. Bacterium m,esenteroides Migula. (Ba- cillus No. XVII, Adametz, Landw. Jahrb., 18, 1889, 249; Migula, Syst. d. Bakt., 2, 1900, 312; Bacterium viscosum Chester, Man. Determ. Bact., 1901, 194.) From cheese. Bacterium modestum Bersteyn. (Arb. bakt. Inst. Karlsruhe, 3, 1903, 95.) From soil. Bacterium monstrosum Henrici. (Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 47.) From Swiss cheese. Bacterium nephritidis Migula. {Ba- cillus nephritidis inter stitialis Letzerich, Ztschr. f . klin. Med., 13, 188-, 33 ; Migula, Syst. d. Bakt., 2, 1900, 310; not Bac- terium nephritidis Chester, Man. De- term. Bact., 1901, 145.) From urine in cases of nephritis. Bacterium ochraceum Migula. {Ba- cillus viscosus ochraceus Freund, Martin, FAMILY BACILLACEAE 761 Inaug. Diss., Erlangen, 1893, 37; Migula, Syst. d. Bakt., 2, 1900, 333.) From the oral cavity. Bacterium olivae Montemartini. (Atti Inst. Bot. Pavia Univ., 2 ser., 14, 1914, 154.) From diseased olive branches. Bacterium paludosum McBeth. (Soil Sci., 1, 1916, 463.) Filter paper reduced to a white pulp-mass. From two soils in California. Bacterium perittomaticum Burchard. (Arb, bakt. Inst. Karlsruhe, 2, 1898, 11.) Similar to or identical with Bacillus ruminatus (Gottheil, Cent. f. Bakt., II Abt., 7, 1901, 485). From soil. Bacterium pituitans Burchard. (In- aug. Diss., 1897; Arb. bakt. Inst. Karls- ruhe, 2, Heft 1, 1898, 8.) From a brown concretion in a cooked egg. Bacterium, plicativum Weiss. (Weiss, Arb. bakt. Inst. Karlsruhe, 2, Heft 3, 1902, 223; not Bacterium plicativum Migula, Syst. d. Bakt., 2, 1900, v and 453.) From fermenting beets and malt. Bacterium plicatum Henrici. (Hen- rici, Arb. bakt. Inst. Karlsruhe, 1, Heft 1, 1894, 49 ; not Bacterium plicatum Chester, Man. Determ. Bact., 1901, 166.) From brick cheese. Bacterium, pseudaceti Migula. (Ba- cillus No. XV, Adametz, Landw. Jahrb., 18, 1889, 247; Migula, Syst. d. Bakt., 2, 1900, 320; Bacterium turgidum Chester, Man. Determ. Bact., 1901, 195.) From cheese. Characteristic involution forms very similar to those of Bacillus aceti Hansen. Bacterium pseudomycoides Migula. (Migula, Syst. d. Bakt., 2, 1900, 486; Bacillus pseudomycoides roseus Nepveux, These, Fac. Pharm., Nancy, 1920, 112.) From soil. Bacterium pseudovermiculosum Saito. (Jour. Coll. Sci., Imp. Univ., Tokyo, 23, Art. 15, 1908, 62.) Isolated twice from garden air. Bacterium pyrenei No . 1 , No . 2 and No . 3, Metalnikov, Ermolaev and Skobaltzyn. (Internat. Corn Borer Invest., 3, 1930, 28 and Ann. Inst. Past., 46, 1931, 467, 468 and 469 respectively; presumably the same as Bacillus pirenei Pospelov, Lenin Acad. Agr. Sci. (U.S.S.R.), Ann. Rept. 1936, 318-321.) No. 1 from diseased larvae of the corn borer (Pyrausta nubi- lalis) that had become black after death ; No. 2 from larvae that had become brown ; and No. 3 from larvae that had become pinkish-brown. Bacterium radiatum Kern. (Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 438; Bacterium barbatum Migula, Syst. d. Bakt., 2, 1900, 317.) From the stomach of a finch. Bacterium rusticum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 440.) From the stomach of a sparrow. Bacterium sempervivum Migula. (No. XII, Flugge, Ztschr. f. Hyg., 17, 1894, 296; Bacillus lactis No. XII, Kruse, in Flugge, Die Mikroorganismen, 3 Aufi., 2, 1896, 269; Migula, Syst. d. Bakt., 2, 1900,321.) From milk. Bacterium serratum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 451.) From the intestine of a dove. Bacterium sewerini Migula . (Sewerin, Cent. f. Bakt., II Abt., 3, 1897, 709; Migula, Syst. d. Bakt., 2, 1900, 330.) From manure. Bacterium spissum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 446.) From the intestine of a bird. Bacterium sp uticola Migula. (Bacillus No. 4, Pansini, Arch. f. path. Anat., 122, 1890, 440; Migula, Syst. d. Bakt., 2, 1900, 306 ; Bacterium sputi Chester, Man. Determ. Bact., 1901, 190.) From sputum. Bacterium streptococciforme Migula. {Bacillus thermophilus III, Rabino- witsch, Ztschr. f. Hyg., 20, 1895, 156; Migula, Syst. d. Bakt., 2, 1900, 343; Bacterium therm,ophilum III, Chester, Man. Determ. Bact., 1901, 185.) From soil, feces, corn. Bacterium subdenticulatum Migula. {Bacillus thermophilus VII, Rabino- witsch, Ztschr. f. Hyg., 20, 1895, 158; Migula, Syst. d. Bakt., 2, 1900, 345; Bac- terium thermophilum VII, Chester, Man. Determ. Bact., 1901, 185.) From feces. 762 MANUAL OF DETERMINATIVE BACTERIOLOGY Bacterium subrubeum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 450; Bacillus snbrubeus Nepveux, These, Fac. Pharm., Nancy, 1920, 115.) From the intestines of birds. Bacterium subsquamosum Migiila. {Bacterium squamosum longum Kern, Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 458; Migula, Syst. d. Bakt., 2, 1900, 335.) From the intestines of a dove. Bacterium subthermophilum Migula. {Bacillus thermophilus IV, Rabino- witsch, Ztschr. f. Hyg., 20, 1895, 157; Migula, Syst. d. Bakt., 2, 1900, 344; Bacterium thermophilum IV, Chester, Man. Determ. Bact., 1901, 186.) From soil and feces. Bacterium subtilis var. galleriae Chorine. (Ann. Inst. Past., 41, 1927, 1120.) From diseased larvae of the bee moth {Galleria mellonella). Bacterium tenax Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 443.) From the stomachs of birds. Bacterium terrae (Ucke) Chester. {Streptobacillus terrae Ucke, Cent. f. Bakt., I Abt., 23, 1898, 1001; Chester, Man. Determ. Bact., 1901, 199.) From soil. Bacterium truncatum Chester. (Ba- cillus No. 51, Conn, Storrs Agr. Exp. Sta., 1894, 81; Chester, Man. Determ. Bact., 1901, 195; not Bacterium trunca- tum Migula, Syst. d. Bakt., 2, 1900, 407; not Bacterium truncatum. Chester, ibid., 157.) From milk. Bacterium verrucosum Kern. (Arb. bakt. Inst. Karlsruhe, 1, Heft 4, 1896, 434.) From the stomachs and intestines of birds. Bacterium virgula (Duclaux) Migula. {Tyrothrix virgula Duclaux, Ann. Inst. Nat. Agron., 4, 1882, 23; Migula, Syst. d. Bakt., 2, 1900, 323.) From cheese. Bacterium viride van Tieghem. (Van Tieghem, Bull. Soc. Bot. France, 1880, 174; Bacilhis viridis Trevisan, I generi e le specie delle Batteriacee, 1889, 18.) Cellulobacillus mucosus Simola. (Ann. Ac. Sc. Fenn., Ser. A, 34, 1931; abst. in Cent. f. Bakt., II Abt., 86, 1932, 89.) Thermophilic ; cellulose decomposed quickly at 55° to 60°C, more slowly at 37°C. Cellulobacillus myxogenes Simola {loc. cit.). Not slimy as above. Clostridium gelatinosum Laxa. (Eine thermophilen Bacillus, Laxa, Cent. f. Bakt., II Abt., 4, 1898, 362; Laxa, ibid., 6, 1900, 286; 8, 1902, 154; Bacterium sacchariphilum Migula, Syst. d. Bakt., 2, 1900, 341 ; Bacterium laxae Chester, Man. Determ. Bact., 1901, 187.) From sugar factory wastes. Produces slime in sucrose solutions. Probably a variety of Bacillus vulgatus according to Sacchetti (Cent. f. Bakt., II Abt., 95, 1936, 115). Denitrobacterium thermophilum Am- broz. (Cent. f. Bakt., II Abt., 37, 1913, 3.) From soil. Lactobacillus sporogenes Horowitz- Wlassowa and Nowotelnow. (Cent. f. Bakt., II Abt., 87, 1933, 331.) Resem- bles Lactobacillus delbrueckii but forms ellipsoidal, terminal spores. M etabacterium polyspora Chatton and Perard. (Compt. rend. Soc. Biol., Paris, 65, 1913, 1232.) The type species of the genus M etabacterium, characterized by forming one to eight spores in a single cell. From the caecum of guinea pigs. See Buchanan (Jour. Bact., 3, 1918, 39). Myxobacillus betae Gonnermann. (Oestcrreich-Ungarische Ztschr. f. Zuck- crind. u. Landwirtsch., 36, 1907, 877; see Cent. f. Bakt., II Abt., 21, 1908, 258.) Produces slime in sucrose solutions. Appears to be closely related to Bacillus subtilis. Nilrosobacillus thermophilus Campbell. (Science, 75, 1932, 23.) A thermophilic aerobic rod with swollen clavate sporan- gia; forms nitrites from ammonium salts. From surface layers of soil from North Carolina and Florida. Semiclostridium commune, S. citreum, S. flavum and S. rubrum Maassen. (Arbt. a. d. biol. Abt. f. Land- u. Forst- wirtsch. am kaiserl. Ges. Amt., 5, 1907, 1.) Produce slime in sucrose solutions. FAMILY BACILLACEAE 763 Germs II. Clostridium Prazmowski* (Prazmowski , Untersuchungen Uber die Entwickelungsgeschiehte und Ferment- wirkung einiger Bacterien-Arten, Inaug. Diss., Leipzig, 1880, 23; Vibrio Mtiller, Vermium terrestrium et fluviatilum, 1, 1773, 39; Bacterium Ehrenberg, Evertebrata, Berlin, 1828, (8?)**; Metallacter Perty, Zur Kenntniss kleinster Lebensformen, 1852, (180?); Amylohacter Trecul, Compt. rend. Acad. Sci., Paris, 61, 1865, 435; Bacillus Cohn, Beitr. z. Biol. d. Pflanzen, /, Heft 2, 1872, 175; Tyrothrix Duclaux, Ann. Inst. Nat. Agron., J^, 1882, (79?); Pacinia Trevisan, Atti della Accad. Fis. -Med. -Statist., Milano, Ser. 4, 3, 1885, (83?) ; Cornilia Trevisan, I generi e le specie delle Batteriacee, Milano, 1889, 21; Gramilohacter Beijerinck, Verhandl. d. k. Akad. v. Wetensch., Amsterdam, Tweedie Sect., Deel I, 1893,4; Bactridium, Paraplectrum, Diplectridiuin and Plectridium Fischer, Jahrb. f. Wissensch. Botan., 27, 1895, 139; Granulobacillus Schattenfroh and Grassberger, Cent. f. Bakt., II Abt., 5, 1899, 702; Streptobacillus Rist and Khoury, Ann. Inst. Past., 16, 1902, 70; Botxdobacillus, Butyribacillus, Cel- lidobacillus , Putribacillus and Pectobacillus Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 342-343; Pectinobacter Makrinov, Arch. Sci. Biol. (Russ.), 18, 1915, 442; Bac- teroides Castellani and Chahners, Man. of Trop. Med., 3rd ed., 1919, 959; Butyriclos- tridium and Putriclostridium Orla-Jensen, Jour. Bact., 6, 1921, 263; Rivoltillus and Metchnikovillus Heller, Jour. Bact., 6, 1921, 550; Omelianskillus, Macintoshillus, Douglasillus, Henrillus, Flemingillus , Vallorillus, Multifermentans, Hiblerillus, Welchillus, Stoddardillus , Arloingillus, Meyerillus, Novillus, Seguinillus, Reglillus, Robertsonillus , Nicollaierillus, Mariellillus, Recordillus, Tissierillus, Putrificus, Ermengemillus and Weinbergillus Heller, Jour. Bact., 7, 1922, 5-9; Peptoclosiridium Donker, Inaug. Diss., Delft, 1926, 23 ; Botulinus, Chauvoea and Welchia Pribram, Jour. Bact., 18, 1929, 374; Anaerobacillus , Verrucosus and Euclostridium Janke, Cent. f. Bakt., II Abt., 80, 1930,490; Butylobacter Bakonyi, U. S. Letters Pat., 1,818,782, 1931 ; Caduceus, Endosporus, Infiabilis, Palmula and Terminosporus Pr^vot, Ann. Inst. Past., 61, 1938, 76-86; Acuformis (syn. Palmula) Prevot, Man.d. Class., etc., 1940, 152.) From Greek, Clostridium, a little spindle. Rods, frequently enlarged at sporulation, producing clostridial or plectridial forms. The cells possess no catalase. Anaerobic or microaerophilic. Biochemically very active. Many species ferment carbohydrates producing various acids (frequently including butyric) and gas (CO2, Ho and sometimes CH4). Others cause rapid putre- faction of proteins producing offensive odors. Commonly found in soil and in human or animal feces. Some species, while growing saprophytically on decomposing vege- table matter or on dead tissue within an animal host, form various toxic and lytic substances and are thereby pathogenic. The type species is Clostridium butyricum Prazmowski. Key to the species of genus Clostridium. I. Strictly anaerobic. A. Not typically fermenters of cellulose. 1. Do not characteristically produce distinctive pigments, a. Spores central, excentric, to subterminal. b. Spores oval. * Revised by Prof. R. S. Spray, School of Medicine, West Virginia University, Morgantown, West Virginia, November, 1938; further revision May, 1942. ** In a few instances the original records were inaccessible. In such cases the page is indicated as (8?). In all other cases the page indicates what is believed to be the earliest record of the designation cited. 764 MANUAL OF DETERMINATIVE BACTERIOLOGY c. Rods distinctly swollen at sporulation. d. Motile. e. Gelatin, or glucose gelatin, not liquefied, f. Glucose fermented, g. Coagulated albumin not liquefied. h. Stormy fermentation, or at least active coagulation of milk. Also see hhh. i. Glycerol not fermented, j. Mannitol fermented. k. Starch, lactose and sucrose fermented. 1. Clostridium butyricum. kk. Starch not fermented. Lactose and sucrose fermented, la. Clostridium beijerinckii. jj. Mannitol not fermented. k. Starch and lactose not fermented, lb. Clostridium pasteurianum. ii. Glycerol fermented, j. Mannitol not fermented. k. Starch, lactose, sucrose and salicin fermented. Ic. Clostridium multifermentans . hh. Milk slowly coagulated; not stormily. Also see hhh. i. Glycerol and mannitol not fermented. 2. Clostridium, fallnx. ii. Glycerol not recorded. j. Acid, but no gas, from lactose and sucrose. 3. Clostridiuvi fissum. hhh. Milk not coagulated. i. Glycerol not fermented. 4. Clostridium difficile. gg. Coagulated albumin not recorded. h. Milk acidified, but not coagulated. 5. Clostridium, viscifaciens . ee. Gelatin, or glucose gelatin, liquefied. f. Glucose fermented, g. Coagulated albumin not liquefied. h. Milk slowly coagulated. Clot not digested. i. Glycerol and mannitol not fermented, j. Lactose fermented. k. Sucrose not fermented. Salicin fermented. 6. Clostridium septicum. kk. Sucrose fermented. Salicin not fermented. 7. Clostridium feseri. ii. Glycerol fermented. 8. Clostridium hernolyticurn. hh. Milk acidified but not coagulated. i. Glycerol fermented. j. Mannitol not fermented. k. Starch fermented. Lactose, sucrose and salicin FAMILY BACILLACEAE 765 not fermented. Exotoxin formed; toxic on injection but not on feeding. 9. Clostridium novyi. kk. Starch not recorded. 1. Lactose, sucrose and salicin not fermented, m. Adonitol fermented. 10. Clostridium botulinum. mm. Adonitol not fermented. 10a. Clostridium botulinum. Type C. gg. Coagulated albumin slowly to rapidly liquefied. h. Stormy fermentation, or at least active coagulation of milk. Clot not digested. 11. Clostridium acetobutylicum. hh. Milk slowly and softly coagulated ; not stormily. Clot slowly to rapidly digested. i. Glycerol and mannitol not fermented. Also see iii. j. Starch not recorded, k. Lactose fermented. 12. Clostridium aerofoetidum. kk. Lactose not fermented. 13. Clostridium sporogenes. 13a. Clostridium sporogenes var. A. P. Marie. 13b. Clostridium sporogenes var. equine. 13c. Clostridium tyrosinogenes. 13d. Clostridium flabelliferum. 13e. Clostridium parasporogenes. ii. Glycerol fermented. Also see iii. j. Mannitol not fermented. 14. Clostridium parabotulinum Types A and B. iii. Glycerol not recorded. j. Mannitol and starch not recorded. k. Lactose and sucrose weakly fermented. I. Gas formed from carbohydrates. 15. Clostridium saccharolyticum. II. Gas not formed from carbohydrates. 16. Clostridium regulare. ff. Glucose not fermented. (Carbohydrates not fermented.) g. Coagulated albumin not digested. Lab-coagulation of milk; increasing alkalinity. Clot digested. 17. Clostridium hastiforme. gg. Coagulated albumin not recorded. Slow, mildly acid coagulation of milk. Clot digested. 18. Clostridium subterminale. dd. Non-motile. e. Gelatin, or glucose gelatin, not liquefied. 19. Clostridium malenominatum. c. Rods not swollen at sporulation. 766 MANUAL OF DETERMINATIVE BACTERIOLOGY d. Motile. e. Gelatin, or glucose gelatin, liquefied, f. Glucose fermented, g. Coagulated albumin liquefied. Milk slowly coagulated. Clot slowly digested. 20. Clostridium bifermentans. gg. Coagulated albumin not recorded. h. Milk slowly coagulated; slimy, i. Gas formed from glucose. 21. Clostridium mucosum. ii. Acid but no gas from glucose. 22. Clostridium pruchii. ee. Iron-gelatin (Spray), no growth. 23. Clostridium cylindrosporum. dd. Non-motile. e. Gelatin, or glucose gelatin, liquefied, f. Glucose fermented, g. Coagulated albumin not liquefied. h. Milk stormily fermented. Clot not digested, i. Glycerol fermentation variable, j. Mannitol not fermented. Starch, lactose and su- crose fermented. Salicin rarely fermented. Types identified by specific toxin-antitoxin neutralization. 24. Clostridium perfringens Types A, B, C and D. bb. Spores spherical. c. Rods distinctly swollen at sporulation. d. Motile. e. Gelatin, or glucose gelatin, not liquefied, f. Glucose fermented, g. Coagulated albumin not liquefied. h. Milk acidified; slowly and softly coagulated; not stormily. Clot not digested. 25. Clostridium sphenoides. hh. Milk acidified but not coagulated. 26. Clostridium innominatum. cc. Rods not swollen at sporulation. d. Non-motile. e. Gelatin, or glucose gelatin, not liquefied, f. Glucose fermented, g. Coagulated albumin not liquefied, h. Milk acidified but not coagulated. 27. Clostridium filiforme. aa. Spores terminal. b. Spores distinctly oval to ellipsoid, c. Rods distinctly swollen at sporulation. d. Motile. e. (5elatin, or glucose gelatin, not liquefied. Also see eee. f. Glucose fermented. FAMILY BACILLACEAE 767 g. Coagulated albumin not liquefied. h. Milk slowly coagulated. Clot not digested, i. Glycerol not fermented. j. Mannitol fermented. 28. Clostridium sartagoformum. jj. Mannitol not fermented. 29. Clostridium paraputrificum. ff. Glucose not fermented. g. Coagulated albumin not liquefied. Milk unchanged. 30. Clostridium, cochlearium. gg. Coagulated albumin not recorded. h. Milk, or iron-milk (Spray), no growth, i. Carbohydrates not fermented. j. Ethyl alcohol fermented chiefly to caproic acid. 31. Clostridium kluyverii. jj. Ethyl alcohol not fermented to caproic acid. 32. Clostridium acidiurici. ee. Gelatin, or glucose gelatin, liquefied. Also see eee. f. Glucose fermented, g. Coagulated albumin liquefied. h. Milk often, but not always, coagulated. Clot, if formed, not digested. 33. Clostridium capitovale. hh. Milk acidified but not coagulated. Slow peptoniza- tion, i. Glycerol and mannitol not recorded, j. Starch not fermented. 34. Clostridium parabifermentans. jj. Starch not recorded. Lactose weakly fermented. 35. Clostridium ovalare. eee. Gelatin, or glucose gelatin, not recorded. Glucose fermented with acid but no gas. 36. Clostridium zoogleicum. bb. Spores spherical, or nearly so. c. Rods distinctly swollen at sporulation. d. Motile. e. Gelatin, or glucose gelatin, not liquefied. Also see eee. f. Glucose fermented. g. Coagulated albumin not liquefied. h. Milk slowly coagulated, not stormily. Clot not diges- ted. Also see hhh. 37. Clostridium thermosaccharolyticum. hh. Milk not coagulated; unchanged. Also see hhh. 38. Clostridium caloritolerans . hhh. Milk slowly alkalinized; casein slowly separated. 39. Clostridium tetanoides. ee. Gelatin, or glucose gelatin, liquefied. Also see eee. f. Glucose not fermented. 7CS MANUAL OP DETERMINATIVE BACTERIOLOGY g. Coagulated albumin slowly liquefied. h. Milkmay show soft lab-coagulation. Clot not definitely digested. 40. Clostridium tetani. hh. Milk shows slow, soft lab-coagulation. Clot slowly digested. 41. Clostridium lentoputrescens . ff. Glucose wealdy fermented. g. Coagulated albumin slowly liquefied. h. Milk variably coagulated. Clot, if formed, variably digested. 42. Clostridium filamentosum,. eee. Gelatin records at variance. f. Glucose fermented, g. Coagulated albumin not liquefied, h. Milk not coagulated; unchanged. 43. Clostridium tetanomorphum. dd. Non-motile. e. Gelatin, or glucose gelatin, not liquefied, f. Glucose fermented, g. Coagulated albumin not recorded. 44. Clostridium alcaligenes. ee. Gelatin,or glucose gelatin, liquefied. f. Glucose fermented, g. Coagulated albumin not liquefied. 45. Clostridium angulosum. gg. Coagulated albumin liquefied. 46. Clostridium putrefaciens. 2. Characteristically produce pigments of varied colors. a. Spores central, excentric, to subterminal. b. Spores oval. c. Rods distinctly swollen at sporulation. d. Motile. e. Gelatin, or glucose gelatin, not liquefied. f . Black pigment formed around colonies in deep agar. 47. Clostridium nigrificans. ff. Violet pigment formed in potato mash. g. Indole is formed. 48. Clostridium helfantii. gg. Indole is not formed. 48a. Clostridium maggiorai. fff . Green pigment formed on potato slant, g. Indole is formed. 48b. Clostridium derossii. 48c. Clostridium ottolenghii. 48d. Clostridium paglianii. gg. Indole is not formed. 48e. Clostridium lustigii. 48f. Clostridium sclavoi. FAMILY BACILLACEAE 769 ffff. Red pigment formed in potato mash, g. Indole not recorded. 49. Clostridium venturelli. ee. Gelatin, or glucose gelatin, liquefied. f. Red to orange-red pigment formed, especially in starchy media, g. Indole is not formed, h. Stormy fermentation of milk. Clot slowly softened. 50. Clostridium roseum. hh. Slow, spongy coagulation of milk. Clot slowly digested. 51. Clostridium chromogenes. ff . Yellow-orange pigment formed in various media, g. Indole is not formed. h. Milk actively coagulated, not stormily. Clot is not digested. 52. Clostridium felsineum. aa. Spores terminal. b. Spores oval. c. Rods distinctly swollen at sporulation. d. Non-motile. e. Gelatin, or glucose getamin, no liquefied, f. Deep red pigment formed on potato slants. 53. Clostridium carbonei. Typical fermenters of cellulose. I. Do not characteristically produce distinctive pigments, a. Spores terminal. b. Spores distinctly oval to ellipsoid. c. Rods distinctly swollen at sporulation. d. Motile, e. Gelatin, or glucose gelatin, liquefied. Ferments a variety of carbohydrates, other than cellulose, after prolonged cul- tivation. 54. Clostridium spumarum. ee. Gelatin, or glucose gelatin, not recorded. Carbohydrates, other than cellulose, not fermented. 55. Clostridium, werneri. bb. Spores spherical, or nearly so. c. Rods distinctly swollen at sporulation. d. Non-motile. 56. Clostridium cellulosolvens . Characteristic pigments produced in certain media. a. Spores terminal. b. Spores distinctly oval to ellipsoid. Rods distinctly swollen at sporulation. 57. Clostridium dissolvens. bb. Spores spherical, or nearly so. Rods distinctly swollen at sporula- tion. 58. Clostridium omelianskii. 770 MANUAL OF DETERMINATIVE BACTERIOLOGY II. Microaerophilic. Grow customarily as anaerobes, but are able to produce scan sometimes atypical, growth on aerobic agar slants. A. Not typically fermenters of cellulose. 1. Do not characteristically produce distinctive pigments. a. Spores central, excentric, to subterminal. Spores oval. Rods dig tinctly swollen at sporulation. 59. Clostridium carnis. ee. Gelatin, or glucose gelatin, liquefied. f . Carbohydrates not fermented. 60. Clostridium, histolyticum. aa. Spores terminal. Spores distinctly oval to ellipsoid. Rods distinctly swollen at sporulation. 61. Clostridium tertium. 1 . Clostridium butyricum Prazmowski . (Untersuch. ii. d. Entwickelungsgesch- ischte ii. Fermentwirk. einiger Bacterien- Arten, Inaug. Diss., Leipzig, 1880, 23; Bacillus butyricus Fliigge, Die Mikroorg., 2 Aufl., 1886, 296.) From M. L., acidum butyricum, butyric acid. Described from the original incom- plete records of Prazmowski, as amplified by the studies of Adamson, Jour. Path, and Bact., 22, 1919, 371, and of Hall, Jour. Inf. Dis., SO, 1922,467. Rods : 0.7 by 5.0 to 7.0 microns, straight or slightly curved, with rounded ends, occurring singly, in pairs, in short chains and occasional long filaments. Motile. Spores oval, excentric to subterminal, swelling rods to clostridial forms. Gram- positive, becoming Gram-negative. Granulose positive in clostridial stage (blue color with iodine). Gelatin and glucose gelatin : Not liquefied. Plain agar slant (anaerobic) : Little or no growth. Glucose agar surface colonies (anaero- bic): Circular or slightly irregular, slightly raised, moist, creamy-white. Deep glucose agar colonies : Biconvex, dense, yellowish-white, entire. Agar fragmented early by abundant gas. Blood agar not hemolyzed. Plain broth : Little or no growth. Glucose broth : Abundant, diffuse tur- bidity; much gas. Litmus milk: Acid and early coagula- tion. Litmus is reduced. Stormy fer- mentation; clot fragmented but not digested. Indole not formed. Nitrites not produced from nitrates. Fixes atmospheric nitrogen. Acid and gas from xylose, glucose, lac- tose, sucrose, starch, salicin, esculin and mannitol. Amygdalin, pectin, cellulose, glycerol and Ca-lactate not fermented. Fermentation products include butyl, ethyl and iso-propyl alcohols, acetone, organic acids, H2 and CO2. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Non-pathogenic for guinea pig and rabbit. Grows well from 30°C to 37°C. Anaerobic. Source : Originally isolated from cheese. Commonly encountered in naturally soured milk, in naturally fermented starchy plant substances and in soil. Habitat : Probably rather widely dis- persed in soils rich in humus. Note : Many butyric acid-producing anaerobes are recorded in the literature. The questionable purity and the incom- plete descriptions, particularly of the older species, make it difficult to deter- mine the degree of relationship of these species to Clostridium butyricum Praz- mowski. The following list cites the outstanding historic or recently de- scribed species. FAMILY BACILLACEAE 771 Ferment butyrique, Pasteur, Compt. rend. Acad. Sci., Paris, 52, 1861, 345 (Vibrion butyrique, Pasteur, ihid., 1261) ; Bacillus amylobacter van Tieghem, Bull, de la Soc. Botan. de France, U, 1877, 128 {Metallacter amylobacter Trevisan, Reale 1st. Lombardo d. Sci. e Lett., Rendiconti, 1879, 147; Clostridium amylobacter Hol- land, Jour. Bact., 5, 1920, 217); Bac- terium navicula Reinke and Berthold, Untersuch. a. d. Bot. Lab. d. Univ. Gottingen, 1, 1879,21 {Amylobacter navic- ula Wehmer, Cent. f. Bakt., II Abt., 4, 1898, 696; Bacillus navicula Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 10, 1898, 128; Clostridium naviculum Prevot, Ann. Inst. Past., 61, 1938, 80); Bacillus butylicus Fitz, Ber. d. Deuts. Chem. Gesellsch., 15, 1882, 867 {Bac- terium fitz Buchner, Ztschr. f. Physiol. Chem., 9, 1885, 384); Butylbacillus, Buchner, ibid., 391; Clostridium butyr- icum {Bacillus amylobacter) I, II, III, Gruber, Cent. f. Bakt., 1, 1887, 370-371; Bacillus butylicus Migula, Syst. d. Bakt., 2, 1900, 598 {Clostridium butyricum I, Gruber, Cent. f. Bakt., 1, 1887, 370); Bacillus gruberi Migula, loc. cit., 599 {Clostridium butyricum II, Gruber, loc. cit., 371) ; Bacillus subanaerobius Migula, loc. cit., 600 {Clostridium butyricum III, Gruber, loc. cit., 371); Bacille amyl- ozyme, also bacille amylocyme, Perdrix, Ann. Inst. Past., 5, 1891, 290 {Bacillus amylozyma IMigula, Syst. d. Bakt., 2, 1900, 626; Bacillus amylozymicus Peter- son, Scott and Thompson, Biochem. Ztschr., 219, 1930, 1; Clostridium amijl- ozyme Prevot, Ann. Inst. Past., 61, 1938, 79; Clostridium var. amylozyme Prevot, Man. d. Class., etc., 1940, 109); Bacillus orthobutylicus Grimbert, Ann. Inst. Past., 7, 1893, 353; Granulobacier butylicum Beijerinck, Verhaudl. d. K. Akad. V. Wetensch., Amsterdam, Twee- die Sectie, Deel I, 1893, 3 {Clostridium butylicum Donker, Thesis, Delft, 1926, 149; Amylobacter butylicum van Beynum and Pette, Cent. f. Bakt., II Abt., 93, 1935, 200; this species is probably identi- cal with Clostridium butyricum I Gruber, Cent. f. Bakt., I Abt., 1, 1887, 370); Granulobacter saccharobutyricum Bei- jerinck, loc. cit., 3, also in Arch. Xeer- land. d. Sci. Exactes et Nat., 29, 1896, 1 (conamonly identified with Bacillus bu- tylicus Fitz, Ber. d. Deuts. Chem. Gesellsch., 15, 1882, 867; Bacillus humosus Migula, Syst. d. Bakt., 2, 1900, 600; Clostridium saccharobutyricum Don- ker, Thesis, Delft, 1926, 147; Amylo- bacter saccharobutyricus van Beynum and Pette, Cent. f. Bakt., II Abt., 93, 1935, 200); Bacillus saccharobutyricus von Klecki, Cent. f. Bakt., II Abt., 2, 1896, 169; Bactridium butyricum Chudia- kow, Zur Lehre von der Anaerobiose (Russ.), Teil I, 1896, (?), cited by Roth- ert, Cent. f. Bakt., II Abt., 4, 1898, 390; Granulobacillus saccharobutyricus mobilis non-liquefaciens Schattenfroh and Grass - berger. Cent. f. Bakt., II Abt., 5, 1899, 702 (bewegliche Buttersaurebacillus, Grassberger and Schattenfroh, Arch. f. Hyg., 42, 1902, 219; Bacillus saccharo- butyricus 7nobilis Hopffe, Ztschr. In- fekrnkh. d. Haust., 14, 1913, .396; Bacillus amylobacter mobilis Gratz and Vas, Cent, f. Bakt., II Abt., 41, 1914, 509); Plec- tridium pectinovorum Stormer, Mitteil. d. Deuts. Landwirts. Gesellsch., 18, 1903, 195 (Microbe du rouissage, Wino- gradsky, Compt. rend. Acad. Sci., Paris, 121, 1895, 744; Granulobacter pectino- vorum Beijerinck and van Delden, Arch. Neerland. d. Sci. Exactes et Nat., Ser. II, 9, 1904, 423; Clostridium pectinovorum Donker, Thesis, Delft, 1926, U5); Bacil- lus holobulyricus Perdrix, Compt. rend. Soc. Biol., Paris, 57, 1904, 481; Bacillus amylobacter Bredemann, Cent. f. Bakt., II Abt., 23, 1909, 385 {Clostridium amylo- bacter Prevot, Ann. Inst. Past., 61, 1938, IQ) ; Amylobacter nonliquefaciens Rusch- mann and Bavendamm, Cent. f. Bakt., II Abt., 64, 1925, 359; Clostridium inter- medium Donker, Thesis, Delft, 1926, 147 (Strain No. 3, Donker, ibid., 39); Clostridium butyricum iodophilum 772 MANUAL OF DETERMINATIVE BACTERIOLOGY Svartz, Jour. Inf. Dis., 47, 1930, 138 (Clostridium iodophilum Pr^vot, Ann. Inst. Past., 61, 1938, 80); Granulobacter saccharobutyricus immobile nunliquefa- ciens McCoy, Fred, Peterson and Hastings, Jour. Inf. Dis., 46, 1930, 121; Bacillus amylobacter S and W, Wertheim, U. S. Letters Pat., 1,917,676, 1933; Clostridium tyrobutyricum van Beynum and Pette, Cent. f. Bakt., II Abt., 93, 1935, 208; Clostridium polyfermeniicuni, Clostridium saccharopetum, Clostridium saccharophilicum and Clostridium sac- charopostulatum Partansky and Henry, Jour. Bact.. 30, 1935, 564. la. Clostridium beijerinckii Donker. (Donker, Thesis, Delft, 1926, 145.) Named for M. W. Beijerinck, the Dutch bacteriologist. Has the general characters of Clos- tridium hutyricum. Distinctive character : Non-fermenta- tion of starch. Acid and gas from glucose, lactose, sucrose, inulin, galactose, fructose and mannitol. Glycerol and starch not fer- mented. Source : From soil and fermenting plant tissues. Habitat : Apparently widely distrib- uted in agricultural soils. lb. Clostridium pasleurianiun Wino- gradsky. (Winogradsky, Arch. Sci. Biol. (Russ.), 3, 1895, 330; Clostridium pas- torianum Winogradsky, Cent. f. Bakt., II Abt., 9, 1902, 43 ; Bacillus pasteurianus Lehmann and Neumann, Bakt. Diag., 4thAufl.,^, 1907,82; Bacillus pastorianus Lehmann and Neumann, ibid., 462; not Bacillus pastorianus Mac^, Traits Prat, d. Bact., 4th ed., 1901, 957; Bacillus winogradskyi Weinberg et al., Les Mi- crobes Ana^r., 1937, 645.) Named for Louis Pasteur, the French scientist. Probably related species : Bodily, Univ. Colorado Studies, 26, 1938, 30, records 5 new species isolated from 10 strains re- ceived labeled C. pasteurianum. These have been designated as Bacillus dulcito- fermentans, Bacillus rhamnoticus, Bacil- lus inulofugus. Bacillus nonpentosus and Bacillus azoticus. Has the general characters of Clos- Iridium, butyricum. Distinctive characters : Prolonged re- tention of the spore within a peculiar brush-like spore-capsule, and the non- fermentation of starch. Assimilates free atmospheric nitrogen. Distinguished from Clostridium bei- jerinckii by the non-fermentation of lactose and mannitol, and from Clos- tridium, butyricum by the non-fermenta- tion of starch. Acid and gas from glucose, sucrose, inulin, galactose, fructose and dextrin. Glycerol, starch, lactose and mannitol not fermented. Source : Originally isolated from soil . Habitat : Not determined, but appar- ently of restricted and local distribution in soil. Ic. Clostridium multifermentans Ber- gey et al. (Bacillus multifermentans tenalbus Stoddard, Lancet, 1, 1912, 12; Multifermentans tenalbus Heller, Jour. Bact., 7, 1922, 6; Bergey et al., Manual, 1st ed., 1923, 324.) From Latin, multus, many, and fermentans, fermenting. Has the general characters of Clos- tridium butyricum, and is probably only a variety. Distinctive character: Blood agar colonies show a zone of hemolysis in 24 hours. Nitrites are produced from nitrates. Distinguished from Clostridium bu- tyricum by the above characters and by the fermentation of glycerol and non- fermentation of mannitol. Distinguished from Clostridium bei- jerinckii by the fermentation of starch and of glycerol. Distinguished from Clostridium pas- teurianum by fermentation of starch and of lactose. FAMILY BACILLACEAE 773 Acid and gas from glucose, fructose, galactose, maltose, lactose, sucrose, raffi- nose, starch, salicin, inulin and glycerol. Mannitol and dulcitol not fermented. Source : Originally isolated from human gaseous gangrene. Habitat : Found in soil and milk. Widely distributed in nature. 2. Clostridiiun fallax (Weinberg and Seguin) Bergej' et al. (Bacille A, Weinberg and Seguin, Compt. rend. Soc. Biol., Paris, 78, 1915, 277; Bacillus fallax Weinberg and Seguin, ibid., 686; not Bacillus fallax Ornstein, Ztschr. f. Hyg., 91, 1920, 159; Vallorillus fallax Heller, Jour. Bact., 7, 1922, 6; Bergey et al.. Manual, 1st ed., 1923, 325.) From Latin, /aWax, deceptive. Rods : 0.6 by 1.2 to 5.0 microns, occur- ring singly or rarely in pairs. Motile with peritrichous flagella. Encapsulated in body fluids. Spores rarely observed, oval, excentric to subterminal, swelling rods. Gram-positive. Gelatin not liquefied. Glucose agar surface colonies (anaero- bic): Circular, flat, with transparent, crenated margin. Glucose agar deep colonies : Lenticular, bean-shaped, irregular, smooth. Agar slant (anaerobic) : Grayish film. Broth: Poor growth; slight diffuse turbidity. Glucose broth : Abundant turbidity and gas. Clearing by sedimentation. Indole not formed (Duffett, Jour. Bact., 29, 1935, 576). Litmus milk: Acid, slowly coagulated. Litmus reduced. Clot channeled bj- gas, but not digested. Acid and gas from glucose, galactose, fructose, maltose, lactose, sucrose, inu- lin, salicin and starch. Glycerol and mannitol not fermented. Records vary in regard to action on lactose, inulin and salicin. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Meat medium reddened ; not blackened or digested. Pathogenicity for guinea pig variable, and commonlj^ lost in cultivation. Forms a weak exotoxin. Optimum temperature not recorded; grows well at 37°C. Anaerobic. Source: From war wounds, appendici- tis, and once from black-leg of sheep. Habitat : Not determined, other than these sources. 3. Clostridium fissum (Debono) Ber- gey et al. (Bacillus fissus Debono, Cent, f. Bakt., I Abt., Orig., 62, 1912, 232; Bergey et al.. Manual, 1st ed., 1923, 332.) From Latin, fissum, separated. Rods : Variable in size, rounded or square ends, occurring singly, in pairs and in chains and filaments. Motile. Spores small, oval, subterminal, slightly swelling rods. Gram -positive. Gelatin : Not liquefied. Deep gelatin colonies at 22°C : Small, brownish, globular, opaque and entire. Deep glucose agar colonies : Small, white, globular. Gas is formed. No pigment formed. Broth: Uniformly turbid. Milk: Acid, coagulated after 3 days. Indole not formed. Acid and gas from glucose. Acid only in lactose and sucrose. Coagulated albumin not liquefied. Grows at 22°C and 37°C. Anaerobic. Distinctive character : All cultures smell strongly of butyric acid. Source : From human feces. Habitat : Not determined, other than this source. 4. Clostridium difficile (Hall and O 'Toole) Prevot. {Bacillus difficilis Hall and O'Toole, Amer. Jour. Dis. Child., 49, 1935, 390; Clostridium difficilis Pr(5vot, Ann. Inst. Past. 61, 1938, 84.) From Latin, difficilis, difficult. Rods :Heavy-bodied. Actively motile. 774 MANUAL OF DETERMINATIVE BACTERIOLOGY Spores elongate, subterminal slightly swelling rods. Gram-positive. Gelatin : Not liquefied. Blood agar surface colonies (anaerobic) : Irregular, flat and non-hemolytic. Deep agar colonies : Minute, flat, opaque disks, becoming lobate. Milk: Poor growth. Gas formed in traces, but milk unchanged. Acid and gas from glucose, fructose, mannitol, salicin and xylose. Traces of gas, but no acid, from galactose, maltose, sucrose, lactose, raffinose, inulin and glycerol. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium with iron is moderately blackened. Digestion not recorded. Pathogenic for guinea pig and rabbit. Subcutaneous inoculation induces marked edema. Death may occur in from 1 to 9 days. Toxicity: Glucose broth culture fil- trates kill guinea pig and rabbit in 24 to 36 hours. Grows well at 37°C. Anaerobic . Source : From feces of new-born infants . Habitat : Not determined, other than this source. 5. Clostridium viscifaciens Sherman and Erb. (U. S. Pat., 2,017,572, 1935.) From Latin, viscus, birdlime, glue; fa- ciens, making. Rods : Vegetative cells 3 to 10 microns long; average about 6 microns. Motile. Spores oval, 1 by 2 microns, central to subterminal, sometimes swelling rods to club-like and spindle-shaped cells. Gram -negative. Granulose reaction positive. Gelatin : Not liquefied. Plain agar slant (anaerobic ) : No growth. Plain agar stab : No growth. Liquid media: Tendency toward floc- culent growth. Milk: Acidified but not coagulated. Casein not digested. Corn mash : Not fermented or digested. Indole not formed. Nitrites produced from nitrates. Ammonia produced from peptone. Acid, gas and alcohols produced from glucose and maltose. Acid and gas from sucrose, lactose, dextrin, starch, glycerol, mannitol and salicin. Calcium lactate : Not fermented. Fermentation products include butyl alcohol (66 parts), iso-propyl alcohol (31 parts), and small amounts of acetone (3 parts). Limiting reaction for growth : About pH 4.0 to about pH 8.0. Optimum temperature 32°C to 36°C. Grows from 15°C to 42.5°C. Anaerobic. Distinctive character : In fermentable sugar broths it produces a copious floc- culum. Source : From soil and from grains and other plant materials in contact with soil. Habitat : Apparently widely dispersed in agricultural soils. G. Clostridium septicum (Mace) Ford.* (Vibrion septique, Pasteur and Joubert, Compt. rend. Acad. Sci., Paris, 85, 1877, 113, and Bull. Acad. Med., 2° Ser., 6, 1877, 794; Vibrio pasteurii Trevisan, Reale 1st. Lombardo d. Sci. e. Lett., Rendiconti, Ser. 2, 12, 1879, 147; Bacillus septicus Mac6, Traits Prat. d. Bact., 1st ed., 1888, 455; not Bacillus septicus Migula, Syst. d. Bakt., 2, 1900, 646 (Un- named aerobic bacillus, Babes, Sept. Proc. d. Kindesalters, Leipzig, 1889, 32; Bacillus septicus ulceris gangraenosi * Note: In an editorial. Jour. Amer. Vet. Med. Assoc, 62, 1922-23, 565, the name Clostridium septicum is ascribed to Winslow et al.. Jour. Bact., 5, 1920, 191. Search fails to confirm the reference. Casual mention is not regarded as sufficient to establish priority. Hence, Ford is regarded as the author of this binomial. FAMILY BACILLACEAE 775 Sternberg, Man. Bact., 1893, 472); not Bacillus septicus Klein, Micro-organisms and Disease, 1884, 78; Cornilia pasteuri Trevisan, I generi e le specie delle Bat- teriacee, 1889, 22; Bacillus septicus gan- grenae Arloing, Legons sur la tuberculose et certaines septicemics, Paris, 1892, 451 ; Vibriogene septique, Rosenthal, Compt. rend. Soc. Biol., Paris, 64, 1908, 398; Vibrio septique LeBlaye and Guggenheim, Man. Prat. d. Diag. Bact., 1914, 438; Rivoltillus vibrion Heller, Jour. Bact., 7, 1922, 6; Bacillus parasarkophysematos Miessner, Cent. f. Bakt., I Abt., Orig., 89 (Bhft.), 1922, 126, and Deuts. Tierarztl. Wchnschr., 30, 1922, 416 (Bacillus para- sarcophysematos Zeissler, Cent. f. Bakt., 1 Abt., Orig., 89 (Bhft.), 1922, 119); Vibrio septicus Rottgardt, Deuts. Tier- arztl. Wchnschr., 34, 1926, 553; Ford, Textbook of Bact., 1927, 726; Clostridium septicus Scott, Cornell Vet., 18, 1928, 259; Clostridium septique Topley and Wilson, Princ. of Bact. and Immunol., 1st ed., 2, 1929, 1161.) From Greek, septicus, pu- trefactive, septic. Probable s3'nonym : Bacillus of Ghon and Sachs, Cent. f. Bakt., I Abt., Orig., 34, 1903, 289. Identical or closely related species : Clostridium balaenae Prevot, Ann. Inst. Past., 61, 1938, 81 (Walfischseptikamie Bacillus, Nielsen, Cent. f. Bakt., 7, 1890, 269; Bacille de lasepticemie de la baleinc, Christiansen, Compt. rend. Soc. Biol., Paris, 83, 1920, 324; Walfischbazillus, Christiansen, Cent. f. Bakt., I Abt., Orig., 84, 1920, 127); Bacillus gastromy- cosis ovis Kitt, Bakt. u. Path. Mikros., 2 Aufl., 1893, 239 (Bradsotbacillus, Niel- sen, Monats. Prakt. Tierhlk., 8, 1897, 59) ; Bacillus tumefaciens Wilson, Lancet, 196, 1919, 657 (Clostridium tumefaciens Prevot, Ann. Inst. Past., 61, 1938, 81); not Bacillus tumefaciens Israilsky, Cent, f. Bakt., II Abt., 67, 1926, 236; Bacillus sen Clostridium sarcophysematos bovis Kitt, Bakterienkunde u. Path. Mikros., 2 Aufl., 1893, 232 (Bacillus sarcophysematos Kitt, ibid.. Index, X; not Bacillus sar- cophysematos Zeissler, Cent. f. Bakt., I .\bt., Orig., 89 (Bhft.), 1922, 119.) (See Clostridium feseri.) Confused in the older literature with Koch's bacillus of malignant edema, Mitt, a. d. kais. Gesundhts., 1, 1881, 54 (Ba- cillus oedematis maligni Zopf, Die Spalt- pilze, 3 Aufl., 1885, 88; Clostridium oede- matis malignis Fischer, Jahrb. f . Wissen. Bot., 27, 1895, (146?); Bacillus oedematis Schroeter, in Cohn's Kryptogamen-Flora V. Schlesien, 3, 1, 1886, 163; Clostridium edematis Holland, Jour. Bact., 5, 1920, 218; Clostridium oedematis-maligni Ber- gey et al., Manual, 1st ed., 1923, 325). It is commonly believed at present that Koch's bacillus of malignant edema was a culture of Clostridium septicum contami- nated with Clostridium sporogenes or some closely related organism. Described from Weinberg and Seguin, La Gang. Gaz., Paris, 1918, 79, and from Hall, Jour. Inf. Dis., 30, 1922, 486. Rods : 0.6 to 0.8 by 3.0 to 8.0 microns, rounded ends, occurring singly, in pairs and in short chains in cultures; long chains and filaments commonly predomi- nate in body exudates. Motile, with peritrichous flagella. Spores oval, ex- centric to subterminal, swelling rods. Gram-positive. Gelatin: Liquefied, with gas bubbles. Agar surface colonies (anaerobic): Small, transparent, of variable shape. Blood agar surface colonies (anaerobic) : Delicate, fiat, leaf-like, irregular. He- molytic. Deep agar colonies: Variable; usually finely filamentous, cottony, spherical. Broth: Slight, diffuse turbidity, with clearing. Litmus milk: Litmus reduced; slow coagulation and moderate gas. Clot not digested. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, maltose, lactose and salicin. Sucrose, inulin, mannitol and glycsrol not fermented (Hall, loc. cit., 489). Coagulated albumin not liquefied. Blood serum not liquefied. 776 MANUAL OF DETERMINATIVE BACTERIOLOGY Brain medium not. blackened or di- gested. Meat medium reddened; not blackened or digested. Pathogenic lor guinea pig, rabbit, mouse and pigeon. Forms an exotoxin for which an antitoxin is prepared. Optimum temperature about 37"C. Anaerobic. Source : Originallj" isolated from ani- mals inoculated with soil; later from ma- lignant edema of animals, and from human war wounds and from appendicitis. Habitat : Animal intestine, and in manured soils. 7. Clostridium feseri Trevisan. (Be- weglichen Bakterien, Feser, Ztschr. f. Prakt. Vet.-Wissensch., 4, 1876, 19; Tre- visan, Atti Accad. Fis. -Med. -Stat, di Milano, S, 1885, 116; Bacterium chauvoei Arloing, Cornevin and Thomas, Le char- bon symptomatique du boeuf, Paris, 2nd ed., 1887, 82; Bacillus chauvoei De Toni and Trevisan, in Saccardo, Sylloge Fun- gorum, 8, 1889, 1004; Bacillus chauvaei Trevisan, I generi e le specie delle Bat- teriacee, 1889, 22; Bacillus feseri Kitt, Bacterienkunde, etc., 2 Aufl., 1893, 233; Bacillus anthracis symptomatici Kruse, in Fliigge, Die Mikroorg., 3 Aufl., 2, 1896, 245; Bacillus carbonis Migula, in Engler and Prantl, Die nattir. Pfianzenfam., 1, la, 1895, 26; Butyribacillus chauvoei Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 342; Bacillus gangraenae emphy- sematosae Hutyra and Marek, Spez. Path. u. Ther. d. Haust., 3 Aufl., 1, 1910, 39; Bacillus chauvei Holland, Jour. Bact., 5, 1920, 217; Clostridium chauvei Holland, ibid., 217; Bacillus anthracis -sympto- m,atici Holland, ibid., 217; Clostridium anthracis-symptomatici Holland, ibid., 217; Bacillus sarkophysematos Miessner, Cent. f. Bakt., I Abt., Orig., 89 (Bhft.), 1922, 123 {Bacillus sarcophysematos Zeiss- ler, ibid., 119; not Bacillus sarcophysema- tos Kitt, Bakterienkunde, etc., 2 Aufl., 1893, Index, X); Bacillus symptomaticus Matouschek, Cent. f. Bakt., II Abt., 58, 1923, 472; Clostridium chauvoei Scott, Jour. Inf. Dis., 38, 1926, 262; Clostridium chauvaei Scott, Cornell Vet., 18, 1928, 259. ) Named for Feser, an early German bacteriologist. Possible synonyms : Bacterio ocello cuneato, Rivolta, Giorn. di Anat., Fisiol. e Patol. d. Animali, Piso, U, 1882, 33; Bacillum cuneatum, Rivolta, ibid., 67; Bacillum ocello-cuneatum, Rivolta, ibid., 67; Bacteriutn cuneatum Rivolta, ibid., 77; Bacterium ocello cuneatum Rivolta, ibid., 78; Bacillus sarcophysematosi Pep- pier, Cent. f. Bakt., I Abt., 29, 1901, 354. Rods : 1.0 by 3.0 to 8.0 microns, occur- ring singly, in pairs and in short chains. Usually show a dark chromatic point near each end. Motile with peritrichous flagella. Spores oval, excentric to sub- terminal, swelling rods. Gram-positive. Gelatin: Liquefied. Gas bubbles. Agar surface colonies (anaerobic): Small, grayish, semi-opaque, filamentous. Agar slant (anaerobic): Grayish, spreading growth. Broth : Turbid, slightly peptolytic. Litmus milk: Acid; slowly coagulated. Gas maj' be formed. Clot not digested. Indole not formed (early studies record only a trace). Acid and gas from glucose, fructose, galactose, maltose, lactose and sucrose. Inulin, salicin, mannitol, glycerol and dextrin not fermented (Hall, Jour. Inf. Dis., 30, 1922,486). Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or digested. Egg-meat medium : Small gas bubbles in 8 hours. Meat becomes pinkish and the liquid slightly turbid. No blacken- ing or digestion. Pathogenic for guinea pig, mouse and rabbit. Forms an exotoxin. Optimum temperature 37°C. Can grow at 50°C. Anaerobic. Source : The cause of black leg, black quarter or symptomatic anthrax in cattle and other animals. Habitat: Probably soil; especially where heavily manured. FAMILY BACILLACEAE 777 8. Clostridium hemolyticiun (Hall) Hauduroy et al . {Clostridium hemolyticus bovis Vawter and Records, Jour. Amer. Vet. Med. Assoc, 68 (N.S. ^1), 1925-26, 512; Bacillus hemolyticus Hall, Jour. Inf. Dis., 45, 1929, 156; Hauduroy et al.. Diet. d. Bact. Path., 1937, 125.) From Greek, haem^, blood; lyticus, dissolving. Related species : Clostridium hemo- lyticum var. sordelli Hauduroy et al., loc. cit., 126 (Bacillus sp. (?), Sordelli, Prado and Ferrari, Compt. rend. Soc. Biol., Paris, 106, 1931, 142; Unnamed anaerobe of Matte, Inst. Biol. Soc. Nac. Agric.,Chile, ;?A, 1921, (31?) (cited from Vawter and Records, loc. cit., 172). Rods: 1.0 to 1.3 by 3.0 to 5.6 microns, with rounded ends, occurring singly, in pairs and in short chains. Motile with long peritrichous flagella. Spores oval to elongate, subterminal, swelling rods. Gram-positive. Gelatin: Liquefied. Blood agar surface colonies (anaerobic) : Light, diffuse growth. Blood hemolyzed. Deep agar colonies : At first lenticular, becoming densely woolly masses with short peripheral filaments. Little or no gas formed. Broth plus liver : Luxuriant diffuse turbidity, followed by agglutinative clearing. Moderate gas formed. Milk : Acid and slow coagulation. Clot not digested. Acid and gas from glucose, fructose, galactose and glycerol. Lactose, maltose, sucrose, raffinose, arabinose, xylose, inulin, salicin, mannitol and dulcitol not fermented. Subsequent studies show that pure galactose is not fermented (Records and Vawter, Nevada Agr. Exp. Sta., Bull. 173, 1945, 48 pp.). Indole is formed. Methyl red and Voges-Proskauer tests are negative. Nitrites are not produced from nitrates. Hydrogen sulfide is produced. The four characteristics given above are from Records and Vawter (loc. cit., 30). Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or digested. Meat medium reddened, not black- ened. No digestion. Pathogenic and toxic for guinea pig and rabbit. Effect due to an unstable hemo- lytic toxin. Grows well at 37°C. Anaerobic . Source : From blood and tissues of cattle dying of icterohemoglobinuria. Habitat : Not determined. Thus far isolated only from animals. 9. Clostridium novyi (Migula) Bergey et al. (Bacillus oedematis maligni No. II, Novy, Ztschr. f. Hyg., 17, 1894, 212; Bacillus oedematis thermophihis Kruse, in Fltigge, Die Mikruorg., 3 Aufi., 2, 1896, 242; Bacillus novyi Migula, Syst. d. Bakt., 2, 1900, 872; Bacterium oedematis ther- mophilus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 126; Bacillus ther- mophilus Chester, Man. Determ. Bact., 1901, 265; Bacillus oedeviatiens Weinberg and Seguin, Compt. rend. Soc. Biol., Paris, 75, 1915, 507 (Bacille B, Weinberg and Seguin, ibid., 177); Novillus maligni Heller, Jour. Bact., 7, 1922, 7; Clos- tridium oedematiens Bergey et al., Manual, 1st ed., 1923, 326; Bergey et al., idem; Clostridium thermophilum Pribram, Jour. Bact., 22, 1931, 430; Clostridium novyi Type A, Scott, Turner and Vawter, Proc, 12th Internat. Vet. Congr., 2, 1934, 175.) Named for F. G. Novy, the American bacteriologist who first iso- lated this organism. Related or possibly identical species : Neuen pathogenen anaeroben Bacillus, Kerry, Osterr. Ztschr. f. Wiss. Veterin- ark., 5, 1894, 228; Bacterium nivosum LeBlaye and Guggenheim, Man. Prat. d. Diag. Bact., 1914, 344 (Bacille neigeux, Jungano, Compt. rend. Soc. Biol., Paris, 62, 1907, 677; Bacillo nevoso, Jungano, II Tommasi, 2, 1907, (731?); Gasodembazil- lus, Aschoff, Deuts. med. Wchnschr., Jt2, 1916, 512; Bacillus beUonensis Sacqu^p^e, 778 MANUAL OF DETERMINATIVE BACTERIOLOGY Compt. rend. Soc. Biol., Paris, 80, 1917, 850 (Bacille de I'oedeme gazeux malin, Sacquep(5e, ibid., 78, 1915, 316; Clostri- dium bellonensis Prevot, Ann. Inst. Past., 61, 1938, 81); Bacillus gigas Zeiss- ler and Rassfeld, Arch. Wiss. u. Prakt. Tierhlk., 59, 1929, 419 {Clostridium novxji Type B, Scott et al., loc. cit., 175; Clostri- diiim gigas Prevot, Ann. Inst. Past., 61, 1938, 82); not Bacillus gigas Trevisan, Atti. d. Accad. Fis. -Med. -Stat., Milano, Ser. 4, 3, 1885, 96; Clostridium novyi Type C, Scott et al., loc. cit., 175 (non- pathogenic bacillus of osteomyelitis of buffalo, Kraneveld, Nederl. Ind. Bl. Diergeneesk., 4^, 193U, 564; Clostridium bubalorum Prevot, Ann. Inst. Past., 61, 1938, 82; Bacillus osteomyelitis bubalorum Prevot, Man. d. Class., etc., 1940, 123). Rods : 0.8 to 0.9 by 2.5 to 5.0 microns, occurring singly and in pairs, not in chains. Motile with peritrichousflagella. Spores large, oval, subterminal, swelling rods. Gram-positive. Gelatin : Liquefied and blackened. Agar surface colonies (anaerobic): Small, white, with darker center, filamentous. Agar slant (anaerobic): Grayish, spreading growth. Deep agar colonies: Compact, opaque, becoming filamentous with age. Broth: Turbid, with flocculent sediment. Litmus inilk: Acid, not coagulated. Litmus reduced. Acid and gas from glucose, fructose, maltose, xylose, starch and glycerol. Lactose, sucrose, mannitol, dulcitol, inu- lin and salicin not fermented (Hall, Jour. Inf. Dis., 30, 1922,491). Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not lilackened or digested. Pathogenic for guinea pig, rabbit, mouse, rat and pigeon. Forms an exo- toxin, toxic on injection but not on feeding growth. Optimum temperature 35°C to 38°C. Anaerobic. Source : From a guinea pig inoculated with peptonized casein; later from gase- ous gangrene. Habitat : Probably occurs in manured soils. 10. Clostridium botulinxun (Van Ermengem) Holland. {Bacillus botu- linus Van Ermengem, Cent. f. Bakt., I Abt., 19, 1896, 443, and Ztschr. f. Hyg., 26, 1897,48; Holland, Jour. Bact., 5, 1920, 217; Ermengemillus botulinus Heller, Jour. Bact., 7, 1922, 8.) From Latin, botidus, sausage; M.L., botulinus, relating to sausage. Clostridium botulinum comprises a number of toxic species, conveniently divided by Bengtson, U. S. Public Health Serv., Hyg. Lab. Bull. 136, 1924, 33, and by Meyer and Gunnison, Jour. Inf. Dis., 45, 1929, 96 and 108, and by Gunnison and Meyer, Jour. Inf. Dis., 45, 1929, 130, into a non-ovolytic {Clostridium botidinuvi) and an ovolytic {Clostridium parabotuli- nuvi) group. Authorities are not yet in agreement on fermentations and on variant sub-types, and the present group- ings are only tentative, and subject to revision. Meyer and Gunnison cite some 15 sub-types on the basis of toxicity, agglutination and fermentation. The original Van Ermengem strain is not available, and his description is inadequate for classification purposes. Description follows Bengtson {loc. cit.) who used Lister Institute Strain No. 94 (Brit. Med. Res. Counc, Spec. Rept. Ser. No. 12, 1917, 29; ibid.. Spec. Rept. Ser. No. 39, 1919, 26) as a type culture. Rods: 0.5 to 0.8 by 3.0 to 8.0 microns, with rounded ends, occurring singly, in pairs and in short to occasional long chains. Motile with peritrichous flagella. Spores oval, central, subterminal, to terminal at maturation, slightly swelling rods. Gram-positive. Gelatin: Liquefied. Deep liver agar colonies : Fluffy with dense center. Liver agar surface colonies (anaerobic) : No perceptible growth. FAMILY BACILLACEAE 779 Broth: Scant or no growth. Liver broth : Luxuriant turbidity, with considerable gas. Milk: Slowly increasing acidity. No coagulation. No gas. Acid and gas from glucose, fructose, maltose, dextrin, glycerol, adonitol and inositol. Galactose, sucrose, lactose, raffinose, inulin, dulcitol, mannitol, xylose, arabinose, rhamnose and salicin not fermented (Bengtson, loc. cit., 22-25). Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or digested. Meat medium not blackened or digested. Pathogenic for animals. Forms a powerful exotoxin which is neurotoxic both on injection and feeding. Toxin is neutralized by Clostridiuvi parabotu- linum Type B antitoxin. Optimum temperature 20° to 30°C (Van Ermengem, Ztschr. f. Hyg., 26, 1897, 42) ; 30°C (Van Ermengem, Arch. d. Pharmacodyn., 3, 1897, 213 and 499; see Williams and Reed, Jour. Inf. Dis., 71, 1942, 227). Starin (Jour. Inf. Dis., 38, 1926, 103), growth usually earlier at 37°C. Toxin production probably best around 28°C. Anaerobic . Source: Unknown. Culture received through Reddish from Robertson as Bacillus bohilimis No. 94, Strain A, Institute of Infectious Diseases at Berlin. Similar strains have been isolated from canned foods. Habitat : Probably occurs in soil . 10a. Clostridium hotulinum Type C. (Toxin producing anaerobe, Bengtson, U. S. Pub. Health Repts., 37, 1922, 164 and 2252; Bacillus botulinus Type C, Bengt- son, ibid., 38, and U. S. Pub. Health Serv., Hyg. Lab. Bull. 136, 1924, 7; Clostridium luciliae Bergey et al., Man- ual, 1st ed., 1923, 336.) From Latin, botulus, sausage. Probably identical variety : Clostridium parabotidinum equi Theiler and Robin- son, Rev. Gen. de Med. Vet., 36, 1927, 199 {Clostridium boiulinum. Type E, Topley and Wilson, Principles of Bact. and Im- munol., 2nd ed., 1936, 688; Bacillus (Clostridium) botulinum D, Weinberg and Ginsbourg, Donnees Recentes sur les Microbes Anaer., Paris, 1927, 107, but shown to be a Type C by Robinson, Union S. Africa, 16th Ann. Rept., Dir. Vet. Serv. and Animal Indus., 1930, 126; not Clostridium hotulinum Type D, Meyer and Gunnison, vide infra). From a rat carcass presumably responsible for botul- ism in mules in South Africa. Related varieties : Bacillus parabotu- linus Seddon, Jour. Comp. Path, and Therap., 35, 1922, 155 and 275 (Clostri- ditim parabotulinum Ford, Text-Book of Bact., 1927, 743, although this name was used earlier in the "group" sense by Bengtson, U. S. Pub. Health Serv., Hyg. Lab. Bull. 136, 1924, 32). First isolated from bones considered the source of "bulbar paralysis" of cattle in Australia. Clostridium parabotulinus bovis Theiler et al.. Union S. Africa, Dept. Agric, 11th and 12th Repts. of the Dir. Vet. Educ. and Res., Part II, 1927, 1202 (Clostridium botulinum Type D, Meyer and Gunnison, Proc. Soc. Expt. Biol, and Med., 26, 1928-29, 88, also Jour. Inf. Dis., ^5, 1929, 106; not Clostridium botulinum Type D, Weinberg and Ginsbourg, vide supra). From "lamziekte" of cattle in South Africa. Clostridium botulinum Type E, Gunni- son, Cummings and Meyer, Proc. Soc. Expt. Biol, and Med., 35, 1936, 278. An organism received by them from the Russian Ukraine; source of isolation not stated. Clostridium botulinum Type C may be regarded as a variety of Clostridiuvi botulinum, as it has morphologic and cultural characters very similar to those of the Van Ermengem strain. Only divergent or additional characters are recorded here. Rods : 0.5 to 0.8 by 3.0 to 6.0 microns, commonly slightly curved. Agar stab : Slight growth. No gas. 780 MANUAL OF DETERMINATIVE BACTERIOLOGY Deep liver agar colonies : Lenticular, becoming loosely fluffy. Gas is formed. Deep glucose agar colonies: Fluffy, without central nucleus. Gas is not formed. Agar surface growth (anaerobic ) : Very scant, thin. Broth: Scant growth. Milk: Slowly increasing acidity. No coagulation. No digestion. Acid and gas from glucose, fructose, galactose, maltose, glycerol and inositol. Dextrin is weakly fermented. Sucrose, lactose, raffinose, inulin, adonitol, dulci- tol.mannitol, xylose, arabinose, rhamnose andsalicin not fermented. Pathogenic for animals. Forms a powerful exotoxin which is neurotoxic both on injection and feeding. Toxin is neutralized by homologous (Type Ca) antitoxin, but not by Bacillus parabotu- linus Seddon (Type C/3) antitoxin, al- though Seddon-toxin is neutralized by Type Ca antitoxin (Pfenninger, Jour. Inf. Dis.,35, 1924, 347). Grows well at 37°C. Anaerobic. Source: Larvae of blue-bottle fly {Lucilia caesnr). Produces limberneck in chickens. Habitat: Not determined, other than this source. 11. Clostriditun acetobutylicum Mc- Coy, Fred, Peterson and Hastings. (McCoy et al., Jour. Inf. Dis., 39, 1926, 483; ibid., 46, 1930, 118; Clostridium aceto-butylicuvi Legg, U. S. Pat., 1,668,814, 1928; Clostridium acetono- butylicum Pr^vot, Ann. Inst. Past., 61, 1938, 80; Clostridium acetobutyricum Pr^vot, Man. d. Class., etc., 1940, 110.) From Latin, acetum, vinegar and buiyli- cus, butylic, relating to butyl alcohol. Synonyms : Bacillus qranulobacter pectinovoruin Speakman, Jour. Biol. Chem., 41, 1920, 319; Clostridium acetoni- ^enuTODonkcr, Inaug. Diss., Delft., 1926, 144. Rods : Vegetative cells 0.6 to 0.72 by 2.6 to 4.7 microns; Clostridia 1.3 to 1.6 by 4.7 to 5.5 microns. Straight, with rounded ends, occurring singly and in pairs, not in chains. No capsules. Motile with peritrichous flagella. Spores oval , excen- tric to subterminal, swelling rods to Clostridia. Gram-positive, becoming Gram -negative. Granulose reaction positive in clos- tridial stage. Glucose gelatin : Liquefied. Glucose agar surface colonies (anae- robic): Compact, raised, fairly regular. Deep glucose agar colonies : Compact, typically lenticular and smooth. Agar fragmented early by abundant gas. Blood agar not hemolyzed. Pigmentation: None; colonies creamy- white, opaque. Plain broth: No growth. Glucose broth: Abundant, uniform turbidity, with much gas. Litmus milk: Acid and active, often stormy, coagulation. Litmus reduced. Clot fragmented by gas, but not visibly digested. Proteolysis demonstrable, however, on milk agar. Potato : Growth creamy-yellow. Po- tato digested to a yellow slime. Corn mash : Much gas with butylic odor. Indole not formed. Acetylmethylcarbinol formed from many carbohydrates. Nitrites not produced from nitrates. Nitrites reduced to ammonia. Acid and gas from arabinose, xylose, rhamnose, glucose, galactose, mannose, fructose, sucrose, maltose, lactose, raffi- nose, melezitose, starch, dextrin, inulin, glycogen, d-mannitol, a-methyl glucoside and salicin. Esculin, amygdalin and tre- halose are weakly fermented. Melobiose, dulcitol, d-arabitol, perseitol, lactositol, sorbitol, erythritol, adonitol, inositol, quercitol, glycerol, pectin and cellulose are not fermented. Fermentation products include ace- tone, butyl and ethyl alcohols, butyric and acetic acids, H2 and COj. Coagulated albumin cubes : Softened and browned by slow digestion. FAMILY BACILLACEAE 781 Hydrogen sulfide produced from thio- sulfate or sulfite; generally negative from proteinaceous sources. Blood serum not liquefied. Brain medium not blackened or digested. Non-pathogenic for guinea pig and rabbit. Optimum temperature probably about 37°C. Grows from 20°C to 47°C. Anaerobic. Source: From corn, molasses, potato and garden soil. Habitat : Widely, but apparently sparsely, dispersed in agricultural soils. Note : A number of acetone and butyl alcohol-fermenting anaerobes have been described. Present knowledge, however, does not permit any expression of the degree of possible relationship. Only a few well-described species are cited. Bacillus butylicus B. F., Ricard, U. S. Pat., 1,385,888, 1921; Bacillus butylaceti- cum Freiberg, U. S. Pat., 1,537,597, 1925; Clostridium butyricum {Prazmow- ski-Pike-Smyth) Pike and Smyth, U. S. Pat., 1,655,435, 1928; Butylobactcr betae, Butylobacter sinense, Butylobacter solani and Butylobacter zeae Bakonyi, British Pat., 328,723, 1930, and U. S. Pat., 1,818,- 782, 1931; Bacillus saccharobrityricus liqucjaciens McCoy et al.. Jour. Inf. Dis., 46", 1930, 121 (Bacillus saccharobutyricum liquefaciens Legg and Stiles, U. S. Pat., 1, [27 ,813, 1933); Clostridium saccharo- butylicum-gaimna Izsak and Funk, U. S. Pat., 1,908,361, 1933 (Clostridium sac- charobutylicum gamma and Clostridium saccharobutyricum gamma Izsak and Funk, U. S. Pat., 2,016,112, 1935); Clostridium saccharo-acetobutylicum-al- pha McCoy, British Pat., 415,311, 1934; Clostridium propyl-butylicum Muller and Legg, British Pat., 415,312, 1934 (Clostri- dium propyl butylicum Legg and Stiles, U. S. Pat., 2,063,448, 1936); Clostridium saccharobutyl-acetonicum Loughlin, Brit- ish Pat., 409,730, 1934, and U. S. Pat., 1,996,428, 1935, and 1,992,921, 1935; Clos- tridium saccharo-acetobutylicumStilessind Legg, British Pat., 437,121, 1935 (Clostri- dium, saccharo-acetobutylicum Legg, U.S. Pat., 2,063,449, 1936); Clostridium sac- charo-acetobutylicum-beta Arzberger, U. S. Pat., 2,050,219, 1936; Clostridium sac- charo-acetobutylicum-gamma Arzberger, ibid. ; Clostridium inverlo-acetobutylicum Legg and Stiles, British Pat., 437,120, 1935, and Legg, U. S. Pat., 2, 063 ,449, 1936; Clostridium (Bacillus) tetrylium Owen, Mobley and Arroyo, Cent. f. Bakt., II Abt., 95, 1936, 131; Clostridium sac- charobutyl-isopropyl-acetonicum Lough- lin, U. S. Pat., 2,085,666, 1937 (Clostri- dium saccharo-butyl-isopropyl-acetoni- cum Loughlin, ibid., and U. S. Pat., 2,096,377, 1937). 12. Clostridium aerofoetidiim (Wein- berg and Seguin) Bergeyetal. (BacilleD, Weinberg, Compt. rend. Soc. Biol., Paris, 79, 1916, 117; Bacillus aerofoetidus Wein- berg and Seguin, ibid., 1028; Bacillus aero-foetidus Mcintosh, Med. Res. Counc, Spec. Rept. Ser. No. 39, 1919, 42; Seguinillus aerofoetidus Heller, Jour. Bact., 7, 1922, 7; Bergey et al., Manual, 1st ed., 1923, 327.) From Latin, aer, air and foetidus, fetid. Rods : 0.4 to 0.6 by 3.0 to 5.0 microns, occurring singly, iu pairs and in short chains. Motile with peritrichous flagella. Spores rare, oval, subterminal, slightly swelling rods. Gram-positive. Gelatin : Rapidly liquefied. Agar surface colonies (anaerobic) : Circular, transparent, with faintly bluish tint, fimbriate. Deep agar colonies : Lenticular, becom- ing indented and lobate. Blood agar not hemolyzed. Glucose broth : Turbid; with sediment. Litmus milk: Acid; slowly coagulated; followed by slow peptonization. Gas is formed. Acid and gas from glucose, fructose, galactose, mannose, maltose, lactose, xylose, amygdalin, salicin, esculin and glycogen. Sucrose, inulin, glycerol and mannitol not fermented. Coagulated albumin slowly liquefied. Blood serum is liquefied. 782 MAJSrUAL OF DETERMINATIVE BACTERIOLOGY Brain medium blackened and digested. Meat medium reddened, then black- ened and slowly digested. Slightly pathogenic for guinea pig. Optimum temperature 30°C to 35°C. Anaerobic . Source: From gaseous gangrene and from feces . Habitat : Not determined other than these sources. Probably occurs in soil. 13. Clostridium sporogenes (Metchni- koff) Bergeyetal. {Bacillus sporogenes var. A, Metchnikoff, Ann. Inst. Past., 22, 1908, 944; Bergey et al.. Manual, 1st ed., 1923, 329; not Clostridium sporogenes Holland, Jour. Bact., 5, 1920, 220 (Bacil- lus enteritidis sporogenes Klein, Cent. f. Bakt., I Abt., 18, 1895, 737; Bacillus sporogenes Migula, Syst. d. Bakt., 2, 1900, 560; Bacillus {enteritidis) sporo- genes and Bacillus enteritidis Klein, Local Govt. Bd., Ann. Rept. Med. Off., London, 33, 1903-04, 442 and 443.) From Greek, sporus, seed; M.L., spore; genes, producing. Two varieties, A and B, were de- scribed. Bacillus sporogenes var. A, Metchnikoff, loc. cit., 944 {Metchnikovil- lus sporogenes Heller, Jour. Bact., 7, 1922, 9; Clostridium sporogenes var. A, Prdvot, Ann. Inst. Past., 61, 1938, 83) is regarded as the typical form and is de- scribed here. Var. B, see Clostridium bifernientans . Synonyms or probably related species : Oedembacillen, Koch, Mitt. a. d. kaiserl. Gesundheitsamte, 1, 1881, 54; Bacillus oedematis maligni Zopf, Die Spaltpilze, 3 Aufi., 1885, 88 (not Bacillus oedematis maligni Liborius, Ztschr. f. Hyg., 1, 1886, 158; Bacillus oedematis Migula, Syst. d. Bakt., 2, 1900, 604); Bacillus oedematis Chester, Man. Determ. Bact., 1901,292; Clostridium oedematis maligni Bergey et al., Manual 1st ed., 1923, 325 (see Species No. 6, Clostridium septicum Ford, p. 774) : Paraplectrum foetidum Weigmann, Cent, f. Bakt., II Abt., 4, 1898, 827 (Bacterie /3, Weigmann, Cent. f. Bakt., II Abt., 2, 1896, 155; Bacillus weigmanni Chester, Man. Determ. Bact., 1901, 300; Plec- tridium foetidum Weigmann, Mykologie der Milch, Leipzig, 1911, 70; Bacillus anaerobius foetidus LeBlaye and Guggen- heim, Man. Prat. d. Diag. Bact., 1914, 329; Endosporus foetidus Prevot, Ann. Inst. Past., 61, 1938, 75); Bacillus sapro- genes carnis Salus, Arch. f. Hyg., 51, 1904, 114 {Bacillus saprogenes Salus, ibid., 115; not Bacillus saprogenes I, II, III, Herfeldt, Cent. f. Bakt., II Abt., 1, 1895, 77; Bacillus carnis saprogenes Salus, Arch. f. Hyg., 51, 1904, 124; Plectridium saprogenes Pr6vot, Ann. Inst. Past., 61, 1938, 87); Bacillus sporo- genes coagulans Debono, Cent. f. Bakt., I Abt., Orig., 62, 1912, 229 {Clostridium coagulans Bergey et al., Manual, 1st ed., 1923, 335); Reading Bacillus, Don- aldson and Joyce, Lancet, 2, 1917, 448; Bacillus putrifi-cus verrucosus Zeissler, Ztschr. f. Infkrnkh. u. Hyg. Haust., 21, 1920-21, 13 {Bacillus verrucosus Leh- mann and Siissmann, in Lehmann and Neumann, Bakt. Diag., 7 Aufi., 2, 1927, 662). Rods: 0.6 to 0.8 by 3.0 to 7.0 microns, with rounded ends, occurring singly, in pairs, or less frequently in short to long chains and filaments. Motile with peri- trichous fiagella. Spores oval, exccntric to subterminal, swelling rods. Gram- positive. Gelatin: Liquefied and blackened. Agar surface colonies (anaerobic): Small, irregular, transparent, becoming opaque, yellowish-white, fimbriate. Deep agar colonies : Woolly balls with dense, nodular center. Agar slant (anaerobic): Grayish, opaque, spreading. Blood agar is hemolyzed. Broth : Turbid. Gas is formed. Putrid odor. Litmus milk: Softly coagulated. Lit- mus reduced. Slow peptonization, leav- ing a dark, amber-colored liquid. Indole formed (trace). Not formed (Hall, Jour. Inf. Dis., 30, 1922, 482). Nitrites not produced from nitrates. Acid and gas from glucose, fructose, FAMILY BACILLACEAE 783 galactose and maltose. Lactose, sucrose, salicin, glycerol, mannitol and inulin not fermented. (Records vary on many sugars.) Coagulated albumin liquefied. Blood serum liquefied to a dark, putrid liquid. Brain medium blackened and digested. Foul odor. Meat medium reddened, then black- ened and digested with foul odor. Gas is produced. Tyrosin crystals not obvious. Non-pathogenic to guinea pig and rab- bit, other than a slight, temporary local tumefaction. Filtrate non-toxic on in- jection and feeding. Optimum temperature 37°C. Can grow at 50°C. Anaerobic. Source: From intestinal contents, gaseous gangrene, and from soil. Habitat: Common in soil, especially where heavily manured. The following species are commonly regarded as variants of the tj'pical Clostridium sporogenes . 13a. Clostridium sporogenes var. A. P. Marie Pr^vot, Ann. Inst. Past., 61, 1938, 83 (Bacille anaerobie, Marie, Compt. rend. Soc. Biol., Paris, 93, 1925, 21). Resembles the typical Clostridium sporogenes except in the sharp but not putrid odor of its cultures. Pathogenicity : Large abscesses are in- duced on subcutaneous injection into guinea pigs. From spontaneous putrefaction of macerated pork. 13b. Clostridium sporogenes var. equine Pr^vot, Ann. Inst. Past., 61, 1938, 83 (Unnamed anaerobe No. IV, Chouke- vitch, Ann. Inst. Past., 25, 1911, 259). Sporulation is delayed and restricted. Spores are long and almost rectilinear. Litmus milk is coagulated, then the clot is digested after 3 to 4 weeks. Coagulated albumin is slowly dissolved. Not pathogenic for guinea pig or mouse. From large intestine of horse. 13c. Clostridium tyrosinogenes (Hall) Bergey et al. (Culture No. 106, Hall and Finnerud, Proc. Soc. Expt. Biol, and Med., 19, 1921-22, 48; Bacillus tyrosino- genes Hall, Abst. Bact., 6, 1922, 7; not Bacillus tyrosinogenes Rusconi, as cited by Carbone, Ramazotti, Mazzucchi and Monti, Boll. 1st. Sieroter., Milan, 2, 1921, (29?), see Clark and Smith, Jour. Bact., 37, 1939, 278; Bergey et al.. Manual, 1st ed., 1923, 329; Clostridium sporogenes var. tyrosinogenes Pr6vot, Ann. Inst. Past., 61, 1938, 83.) From Greek, tyrus, cheese; M. L., tyrosinum, tyrosine; genes, producing. Ferments monosaccharides but not higher carbohydrates (Hall, Jour. Inf. Dis.,30, 1922,482). Traces of gas, but no acid, from glycerol, sorbitol, mannose, xylose, lac- tose, sucrose, arabinose, galactose, salicin, inulin, dextrin and starch (F. E. Clark, personal communication). Distinctive character : Forms large amounts of tyrosin which precipitate in cultures in protein media. Source : Originally isolated from a cul- ture erroneously labeled Bacillus tetani. Later isolated from an amputated arm. Habitat : Not determined. Only two isolations on record. 13d. Clostridium flabelliferum Sturges and Reddish. (Fish-tailed putrefactive anaerobe. Reddish and Sturges, Abst. Bact., 8, 1924, 5; Sturges and Reddish, Jour. Bact., 11, 1926, 37; Clostridium sporogenes var. caudapiscis Prevot, Ann. Inst. Past., 61, 1938, 83.) From Latin, flabellum, a little fan;/er, bearing. Glucose agar surface colonies (anae- robic): Coarse, raised, with long pe- ripheral intertwining outgrowths. Deep plain agar colonies : Irregular, becoming woolly. Sucrose is fermented (in contrast with Clostridium sporogenes) . Distinctive character : Spores are long 784 MANUAL OF DETERMINATIVE BACTERIOLOGY retained within the sporangium, of which the distal end frays out to fibrils, giving the characteristic fish-tail appearance. Otherwise closely resembles Clostridium sporogenes. Source : From soured hams and from salt. Habitat: Not determined, other than these sources. 13e. Clostridium parasporogenes (Bul- loch et al . ) Bergey et al . (Bacillus Type XII, Mcintosh and Fildes, Med. Res. Counc, Spec. Rept. Ser. No. 12, 1917, 36; Bacillus parasporogenes Bulloch et al., Med. Res. Counc, Spec. Rept. Ser. No. 39, 1919, 39; Bergey et al.. Manual, 1st ed., 1923, 327 ; Clostridium sporogenes var. parasporogenes Pr^vot, Ann. Inst. Past., ei, 1938,83.) Deep agar colonies : Lenticular to slightly irregular. Not woolly. Pathogenic for young guinea pigs. Filtrate non-toxic on injection or on feeding. Optimum temperature 30°C to 35°C. Distinctive character : Resembles Clos- tridium sporogenes, but does not form woolly colonies in deep agar, and is agglutinatively distinct. Probably merely a variety. Source : From gaseous gangrene. Habitat: Not determined. Probably occurs in soil. 14. Clostriditun parabotulinum Bengt- son. (U. S. Pub. Health Serv., Hyg. Lab. Bull. 136, 1924, 32; Types A and B, Burke, Jour. Bact., 4, 1919, 556; Clostri- dium botulinum Types A and B, Bergey et al., Manual, 1st ed., 1923, 328.) From Latin, para, like; M.L., botulinum,, a species name. Note : This group comprises the putre- factive (ovolytic) species, including strains commonly referred to as Memphis and Canton (Type A), and Nevin (Type B). Growth of these types is more easily obtained than with the Clostridium botulinum strains, and the reactions are more obvious. Gunnison and Meyer (Jour. Inf. Dis., 4-5, 1929, 130) propose an intermediate group between Clostridium botulinum and Clostridium parabotulinum, which they call Clostridium metabotulinum. Such a group would provisionally include cer- tain European Type B strains, the Aus- tralian Type C, certain American Type C strains, and the South African Type D. Rods : 0.5 to 0.8 by 3.0 by 8.0 microns, with rounded ends, occurring singly, in pairs, and in short chains. Motile with peritrichous flagella. Spores oval, subterminal, distinctly swelling rods. Gram-positive. Gelatin: Liquehed. Deep liver agar colonies : Type A tend to be restricted to compact disks, with sharp outline and small, opaque nucleus at periphery. Type B tend rather to form loose, woolly colonies (indicative only). Liver agar surface growth (anaerobic) : Profuse, moist. Broth : Fairly abundant diffuse tur- bidity. Many strains spontaneously agglutinate. Liver broth : Luxuriant turbidity. Profuse gas. Milk : Slight acidity ; slow curdling pre- cipitation, with subsequent digestion and darkening. Fermentation records are variable : Acid and gas from glucose, fructose, mal- tose, dextrin, glycerol and salicin. Galactose, sucrose, lactose, rhamnose, raffinose, inulin, adonitol, dulcitol, man- nitol, xylose, arabinose and inositol not fermented (Bengtson, loc. cit., 22-25). Coagulated albumin liquefied: Action of Type B usually more marked than that of Type A. Blood serum liquefied. Brain medium blackened and digested, with putrefactive odor. Meat medium blackened and digested. Putrefactive odor. Tyrosine crystals not observed. Pathogenic for animals. Forms a powerful exotoxin which is neurotoxic both on injection and feeding, and which FAMILY BACILLACEAE 785 is neutralized only by the homologous type antitoxin. Optimum temperature : Records at var- iance. Grows best at 35 to37°C. Toxin production best at about 28°C. Anaerobic . Distinctive character : Types are identified chiefly by protection tests with known-type antitoxin, and to less extent by agglutination. Source: Chiefly from spoiled, non-acid canned goods, from soil and from silage. Habitat : Found rather widely dis- persed in soil. 15. Clostridixun saccharolyticum Ber- gey et al. {Bacillus sporogenes saccharo- lyticus Distaso, Cent. f. Bakt., I Abt., Orig., 59, 1911, 100; Bergeyetal., Manual, 1st ed., 1923, 334.) From Greek, saccha- rum, sugar; lyticus, dissolving, digesting. Rods: Short, rounded ends, occurring singly, in pairs and in short chains. Motile. Spores large, oval, excentric to subterminal, swelling rods. Gram-posi- tive. Gelatin : Liquefied. Deep glucose agar colonies : Gray, bi- convex, lenticular, granular, entire. Gas is formed. Broth: Turbid. Milk: Soft coagulation; casein precipi- tated, then peptonized, leaving a clear, yellow-amber supernatant fluid. Indole is formed. Acid and gas feebly formed from glu- cose. Lactose and sucrose feebly, or doubtfully, fermented. Coagulated albumin slowly liquefied. Grows well at 37°C. Anaerobic . Distinctive character : All cultures give a mixed butyric and fecal odor. Source : From feces of a chimpanzee. Habitat : Not determined, other than this source. 16. Clostridium regulate Bergey et al. {Bacillus sporogenes regularis Distaso, Cent. f. Bakt., I Abt., Orig., 59, 1911, 100; Bergey et al.. Manual, 1st ed., 1923, 334.) From Latin, straight. Long rods : With rounded ends, occur- ring singly and in pairs. Motile. Spores oval, small, subterminal, slightly swelling rods. Gram-positive. Gelatin: Liquefied. Deep agar colonies: Small, opaque, irregular. Milk: Acid; slowly coagulated, then clot slowly digested. Indole formed in small quantity. Slight acidity from glucose, lactose and sucrose. Gas is not formed. Odor of scatol and valerianic acid. Coagulated albumin slowly liquefied. Grows at 37°C. Anaerobic. Source : From human feces. Habitat : Not determined, other than this source. 17. Clostridium hastiforme MacLen- nan. (A4, Cunningham, Cent. f. Bakt., II Abt., 82, 1930-31, 487, and B4a, Cun- ningham, ibid., 83, 1931, 11; MacLennan, Jour. Path, and Bact., 49, 1939, 543.) From Latin, resembling a spear. Rods : Slender, 0.3 to 0.6 by 2.0 to 6.0 microns, with rounded ends, occurring singly, in pairs, and rarely in short chains. Filaments not observed. Motile, with delicate peritrichous flagella; motility persists even after sporulation. Spores ellipsoidal, subterminal, swelling rods. Polar-cap of protoplasm remains long attached to free spores. Gram-positive. Gelatin: Rapidly liquefied. Blacken- ing not recorded. Plain agar surface colonies (anaerobic ) : Minute translucent dots, becoming ir- regularly round, granular, grayish-white, with opaque center and delicate translu- cent border. Blood agar surface colonies (anaerobic) : As above, but larger and more opaque. Old colonies show grayish pigmentation. No hemolysis. Deep plain agar colonies : Small, irregu- larly round with coarsely filamentous r86 MANUAL OF DETERMINATIVE BACTERIOLOGY liorder. A little gas is occasionally formed. Broth: Transient uniform turbidity, quickly settling as a heavy, white, fiocculent deposit. Culture assumes a cheesy odor. Milk: Abundant growth, with lab- coagulation in 2 to 3 days. No increase in acidity; becoming slightly alkaline. Clot completely digested in 10 to 14 days, leav- ing a white, semi-translucent fluid of cheesy odor. Indole not formed. Ammonia not formed. Hydrogen sulfide not formed. Glucose not fermented. Carbohydrates not fermented. Egg medium : Xo digestion or other visible change. Coagulated albumin not digested or blackened. Blood serum not digested or blackened. Meat medium not digested or black- ened, even in presence of metallic iron. Meat particles slightly reddened. Brain medium not digested or blackened. Grows well between 22°(; and 37°C. Anaerobic. Non-pathogenic to guinea pigs on sub- cutaneous inoculation (Cunningham, Cent. f. Bakt., II Abt., 83, 1931, 12). Source : Originally isolated by Cunning- ham as a dissociant from a culture of Bacillus saccharobutyricus von Klecki. Later isolated by MacLennan; 1 strain from a culture of Clostridium, sporogenes , and 2 strains from street dust. Habitat: Not determined, other than these sources. 18. Clostridium subterminale (Hall and Whitehead) comb. nov. {Bacillus subterminalis Hall and Whitehead, Jour. Inf. Dis., 4i, 1927, 66.) Named from the characteristic position of the spores. . Rods : Occurring singly, in pairs and rarely in short chains. Motile. Spores oval, subterminal, swelling rods. Gram- positive. Gelatin: Slowly liquefied, with slight turbidity and black sediment. Blood agar surface colonies (anaerobic) : Delicate. At first mildly, later actively, hemolytic. Deep agar colonies: Opaque, compact, biconvex or lobate discs. Agar slant (anaerobic): No surface growt h . Glucose broth : Turbidity, but no acid or gas formed. Indole not formed. Milk: Slowly coagulated (2 to 3 days), with mild acidity and gas. Slow but com- plete digestion of casein (8 to 18 days). Glucose, fructose, galactose, maltose and lactose not fermented. Brain medium : Slight turbidity in supernatant fluid. Slight gas formation and slow digestion. Iron brain medium : Blackened in 2 to 3 days . Tyrosiu crystals not observable. Non-pathogenic to guinea pigs on sub- cutaneous injection. Grows well at 37°C. Obligately anaerobic. Source: From an African arrowhead. Habitat : Not determined, other than this source. 19. Clostridium malenominatum (Weinberg et al.) comb. nov. (Pseudo-coli anaerobic, Jungano, Compt. rend. Soc. Biol., Paris, 65, 1908-09, 457; Bacillus pseudo-coli anaerobic Jungano and Dis- taso, Les Anadrobies, 1910, 162; Bacillus pseudocoli a7iacrobius LeBlaye and Gug- genheim, Man. Prat. d. Diag. Bact., 1914, 345; Bacillus malenominatus Weinberg et al . , Les Microbes Ana^robies, 1937, 763 ; Paraplectrum malenominatum Prevot, Ann. Inst. Past., 61, 1938, 75.) From Latin, meaning badly named. Rods: Short, cocco-bacillary, becom- ing elongated to short filaments in old cultures — especially in sugar broth. Ends rounded. Distinct bipolar staining tendency. Non-motile. Capsulated, especially in body fluids. Spores oval, FAMILY BACILLACEAE 787 subterminal, slightly swelling rods. Gram-negative. Gelatin : No growth. Deep agar colonies : Small, round, very regular, almost transparent. Gas not formed. Plain broth: Uniform turbidity, set- tling after 48 hours, forming a fine, pow- dery sediment. Indole not produced. jMilk : Growth with no coagulation. Glucose and sucrose not fermented. Coagulated albumin: Not attacked. Meat medium : Abundant growth. No record of changes. Capsules are demon- strable in this medium. Very pathogenic for guinea pigs, which die of septicemia in 24 hours after intra- peritoneal inoculation. Less pathogenic for rabbit, which dies after one week. Toxin not demonstrable in cultures. Grows at 22°C to 37°C. Obligately anaerobic. Source : From feces of a diarrheal infant. Habitat: Not determined, other than this single isolation. 20. Clostridium bifermentans (Wein- berg and Seguin) Bergey et al. {Bacillus bifermentans sporogenes Tissier and Mar- telly, Ann. Inst. Past., 16, 1902, 894; Bacillus hifennantans Weinberg and Seguin, La Gangrene Gazeuse, Paris, 1918, 128; Martellillus bifermentans Hel- ler, Jour. Pact., 7, 1922, 8; Bergey et al.. Manual, 1st ed., 1923, 323.) From Latin, bis, twice, and fermentum, a ferment. Closely related if not identical species : Bacillus centrosporogenes Hall, Jour. Inf. Dis., 30, 1922, 4&4 (Clostridium centro- sporogenes Bergey et al.. Manual, 1st ed., 1923, 322) ; Bacillus oedematis sporogenes Sordelli, Compt. rend. Soc. Biol., Paris, 89, 1923, 55 (Anaerobic agent de gangrene gazeuse, Sordelli, ibid., 87, 1922, 838; Bacillus sordelli Hall and Scott, Jour. Inf. Dis., 41, 1927, 329; Bacillus sporo- genes oedematis Piening, Thesis, Han- over, 1931, (?), cited from McCoy and McClung, The Anaer. Pact., etc., :?, 1939, 492; Clostridium sordelli Prevot, Ann. Inst. Past., 61, 1938, 83); Clostridium oedematoides Meleney, Humphreys and Carp, Proc. Soc. Expt. Biol, and Med., 24, 1926-27, 677. Varying degrees of virulence and toxicity occur in the above group. The more toxic and virulent strains are com- monly referred to as Bacillus sordelli, although otherwise an apparently homo- geneously organized group. Probable synonyms : Clostridium foeti- dum Liborius, Ztschr. f. Hyg., 1, 1886, 160 {Cornilia foetida Trevisan, I generi e le specie delle Batteriacee, 1889, 22; Bacil- lus foetidus Chester, Ann. Rept. Del. Col.Agr. Exp. Sta., 10, 1898, 128; not Bacillus foetidus Trevisan, loc. cit., 16); Bacillus liquefaciens magnus Liideritz, Ztschr. f. Hyg., 5, 1889, 146 {Cornilia magna Trevisan, loc. cit., 22; Bacillus magnus Herfeldt, Cent. f. Bakt., II Abt., 1, 1895, 78; Bacillus magnus lique- faciens LeBlaye and Guggenheim, Man. Prat. d. Diag. Pact., Paris, 1914, 327; Bacillus foetidus clostridiiformis LeBlaye and Guggenheim, idem. 327) ; Clostridium foetidum carnis Salus, Arch. f. Hyg., 51, 1904, 121 {Clostridium carnis foetidum and Clostridium, foetidum Salus, ibid., 121 and 124; Clostridium carnofoetidus McCrudden, Jour. Biol. Chem., 8, 1910- 11, 109; Clostridium carnofoetidum Prevot, Ann. Inst. Past., 61, 1938, 84); Bacillus sporogenes var. B, Metchnikoff, Ann. Inst. Past., 22, 1908, 944 {Clostri- dium sporogenes var. B, Prevot, Ann. Inst. Past., 61, 1938, 83); Bacillus sporo- genes foetidus Chouk^vitch, Ann. Inst. Past., 23, 1911, 257 {Bacillus foetidus Choukevitch, ibid., 258); Bacillus putri- jicus tenuis Zeissler, Ztschr. f. Infkrnkh. u. Hyg. Haust., 21, 1920-21, 13; Bacillus nonfermentans Hall and Whitehead, Jour. Inf. Dis., 41- 1027, 65. Rods : 0.8 to 1.0 by 5.0 to 6.0 microns, occurring singly, in pairs, and in short chains . Spores oval , central to excentric , not distinctly swelling rods. Motile in very young cultures only (less than 24 hours old). Gram-positive. Gelatin : Liquefied and blackened. ■88 MANUAL OF DETERMINATIVE BACTERIOLOGY Agax' surface colonies (anaerobic ) : Circular, crenated to amoeboid. Blood agar surface colonies (anaerobic) : Small, transparent, hemolytic, becoming opaque, yellowish, spreading. Broth : Turbidity and gas. Thick mu- coid sediment. Litmus milk : Slowly coagulated. Slowly peptonized, with little gas. Indole is formed. Nitrites not produced from nitrates. Hydrogen sulfide is produced. Acid and gas from glucose, fructose, mannose and maltose. Lactose, sucrose and inulin not fermented. Records sug- gest variability in glycerol and salicin. Coagulated albumin rapidly liquefied and blackened. Blood serum liquefied and blackened. Brain medium digested and blackened. Egg -meat medium digested and black- ened. Tyrosin crystals in 8 to 10 days. Pathogenicity: Variable with the strain; some kill rabbits in 24 hours; others produce only slight edema, while some show no effect. Toxicity : Likewise variable, from acute to none. Optimum temperature from 30°C to 37°C. Can grow at 50°C. Anaerobic. Source: Originally from putrid meat; subsequently from gaseous gangrene. Habitat: Occurs commonly in feces, soil and sewage. Widely distributed in nature. 21. Clostridium mucosum (Klein) Ber- geyetal. {Bacillus 7)mcosus Klein, Cent, f. Bakt., I Abt., 29, 1901, 991; not Ba- cillus 7nucosus Zimmermann, Die Bakt. unserer Trink- u. Nutzwasser, Chem- nitz, 2, 1894, 8; Bacterium mucosum Migula, Syst. d. Bakt., 2, 1900, 315; Bacillus kleinii Buchanan and Hammer, Iowa Agric. Exp. Sta. Res. Bull. 22, 1915, 276; not Bacillus kleinii Aligula, Syst. d. Bakt., 2, 1900, 766; not Bacillus kleinii Trevisan, in Hit. cited from De Toni and Trevisan, in Saccardo, Sylloge Fun- gorum, 8, 1889, 946; Clostridium kleinii Bergey et al., Manual, 1st ed., 1923, 321; Bergey et al., Manual, 4th ed., 1934, 472; not Clostridium mucosum Simola, cited from Prevot, Man. d. Class., etc., 1940, 112; Endosporus mucosus Prevot, Ann. Inst. Past., 61, 1938, 75.) From Latin, slimy. Rods : 1.3 by 2.0 to 5.0 microns, occur- ring singly, in pairs and in chains. Mo- tile. Spores oval, central, not swelling rods. Gram-negative (Klein, loc. cit.). Young cultures Gram-positive (Buchanan and Hammer, loc. cit.). No growth in media without carbohy- drates. Glucose gelatin: Slowly liquefied. Glucose gelatin surface colonies (anae- robic) : Small, gray. Glucose gelatin stab : Villous growth. Slow liquefaction. Glucose agar slant (anaerobic): Thin, veil-like layer. Slimy condensation water. Glucose broth: Turbid. Gas bubbles. Litmus milk: Acid; slowly coagulated, slimy. Gas formed. Odor of butyric acid. Potato : No growth. Indole not formed. Nitrites not produced from nitrates. Acid and gas from glucose. Blood serum : No growth. Non-pathogenic. Grows at 37°C. Anaerobic. Source: Blood sausage (Blutwurst). Habitat : Not determined, other than this source. 22. Clostriditim pruchii (Buchanan and Hammer) Bergey et al. {Bacillus lactis pruchii Conn, Esten and Stocking, 18th Ann. Rept. Storrs Agric. Exp. Sta., 1906, 179; Bacillus pruchii Buchanan and Hammer, Iowa Agric. Exp. Sta. Res. Bull. No. 22, 1915, 276; Bergey et al.. Manual, 1st ed., 1923, 322.) Named for M. J. Prucha, American bacteriologist. Rods: Variable in size, with club- shaped ends. Motile, with peritrichous FAMILY BACILLACEAE 789 flagella. Spores central, oval, not swell- ing rods. Gram-positive. Gelatin: Rapid, stratiform liquefac- tion. Reddish-yellow sediment. Agar surface colonies (anaerobic) : Round, flat, white, smooth, opaque. Agar slant (anaerobic): Luxuriant, white, viscid. Broth: Turbid, with flocculent pellicle and gray viscous sediment. Litmus milk: Acid; slowly coagulated, becoming slimy yellow. Potato : Thin, brownish, spreading. Indole not formed. Nitrites not produced from nitrates. Acid but no gas from glucose. Coagulated albumin not recorded. Blood serum not liciuefied. Non-pathogenic . Optimum temperature 30°C. Grows well between 20°C and 37°C. Anaerobic. Source : From slimy milk. Habitat : Not determined, other than this source. 23. Clostridium cylindrosporxim Barker and Beck. (Jour. Biol. Chem., m, 1941, 3. ) Named from the characteristic spore morphology. Rods : 1 .0 by 4.0 to 7.0 microns, straight . Motile with peritrichous flagella. Spores elongate to cylindrical, 1.0 to 1.1 by 1.7 to 3.0 microns, central, subterminal to terminal, with little or no swelling of rods. Gram-negative. Iron -gelatin (Spray) : No growth. Deep plain agar : No growth. Deep uric acid agar colonies: Whitish, compact, lobate, 1 to 2 mm in diameter, with irregular edges, surrounded by a zone of precipitated ammonium ureate which gradually disappears. Plain broth: No growth. Glucose broth : No growth. Iron-milk (Spray) : No growth. Indole not recorded (probably nega- tive). Nitrites not recorded (probably nega- tive). Glucose not fermented. Carbohydrates not fermented. Cellulose not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not digested or black- ened. Pathogenicity not recorded (probably non-pathogenic ) . Optimum temperature about 35°C. Optimum reaction about pH 7.5; lower limit for growth pH 6.5. Anaerobic. Distinctive characters : Requires uric acid, or certain other purines, as a pri- mary source of carbon and energy. The purines are converted into ammonia, CO2, acetic acid and a little glycine. This organism is physiologically similar to Clostridium acidi-urici, but may be readily distinguished from the latter by its morphology. Source : A single strain isolated from .soil. Habitat: Probably soil, although only this single isolation is recorded. 24. Clostridium perfringens (Veillon and Zuber) Holland.* Clostridixim ■per- fringens Type A, Wilsdon. {Bacillus aerogenes capsulatus Welch and Nuttall, Johns Hopkins Hosp. Bull. 3, 1892, 81 {Bacillus capsulatus aerogenes Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 327) ; Bacillus phlegmones emphy- sematosae Fraenkel, Ueber Gasphleg- * Because of use of the species name perfringens bj' the Permanent Standards Commission of the Health Organization of the League of Nations (Report of the Per- manent Commission on Biological Standardization, London, June 23, 1931), the use of this name has been continued although it is preceded by a valid binomial {Bacillus emphysematosus Kruse). 790 MANUAL OF DETERMINATIVE BACTERIOLOGY monen, Leipzig, 1893, 47; Bacillus emphysematosus Kruse, in Fliigge, Die Mikroorg., 3 Aufl., 2, 1896, 242; Bac- terium aerogenes capsulatus Chester, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 125; Bacterium emphysematosus Chester, ibid., 126; Bacillus emphy- sematis vaginae Lindenthal, Wien. klin. Wchnschr., 10, 1897, 42; Bacillus per- fringens Veillon and Zuber, Arch. M<5d. Expt. et Anat. Path., 10, 1898, 539; Bacillus capsulatus anaerobius and Ba- cillus capsulatus aerogenes Binaghi, Cent. f. Bakt., II Abt., 4, 1898, 920; Granulo- bacillus saccharobutyricus imvwbilis liquefaciens Schattenfroh and Grass- berger, Cent. f. Bakt., II Abt., 5, 1899, 702 (Granulobacillus immobilis Schatten- froh and Grassberger, Arch. f. Hj^g., 37, 1900, 68; Bacillus amylobacler immobilis Gratz and Vas, Cent. f. Bakt., II Abt., 41, 1914, 509); Bacterium welchii Migula, ibid., 392; Bacillus welchii Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 457; Bacillus butyricus asporogenes im- mobilis Rocchi, Cent. f. Bakt., I Abt., Orig., 60, 1911, 580; probably Bacillus multiformis Distaso, Cent. f. Bakt., I Abt., Orig., 69, 1911, 101 (Bacteroides multiformis Bergey et al., Manual, 1st ed., 1923, 263; Cillobacterium. multiforme Pr^vot, Ann. Inst. Past.. 60, 1938, 297; not Bacillus multiformis van Senus, Inaug. Diss., Leiden, 1890, (?), quoted from Herfeldt, Cent. f. Bakt., II Abt., 1, 1895, 117); Bacillus aerogenes-capsu- latus Holland, Jour. Bact., 5, 1920, 217; Clostridium aerogenes-capsulaturn Hol- land, ibid., 217; Bacillus phlegmoncs- emphysematosae Holland, ibid., 219; Clostridium phlegmones-emphysematosae Holland, ibid., 219; Clostridium phleg- mones emphysematosae Holland, ibid., 222; Clostridium welchii Holland, ibid., 221; Clostridium perfringens Holland, ibid., 219; Welchillus aerogenes Heller, Jour. Bact., 7, 1922, 6; Butyribacillus immobilis-liqucfaciens Heller, ibid., 18; Bacillus welchii Type A Wilsdon, Univ. Cambridge, Inst. Animal Path., 2nd Rept. of Dir., 1931, 72; Clostridium saccharobutyricum liquefaciens van Bey- num and Pette, Cent. f. Bakt., II Abt., 93, 1935-36, 205; Welchia perfringens Pr^vot, Ann. Inst. Past., 61, 1938, 78.) Latinized, very fringed. Related varieties : Clostridium egens Bergey et al., Manual, 1st ed., 1923, 324 {Bacillus egens Stoddard, Jour. Exp. Med., 29, 1919, 187; Stoddardillus egens Heller, Jour. Bact., 7, 1922, Q; Clostridium perfringens var. egens Hauduroy et al., Diet. d. Bact. Path., 1937, 124; Welchia perfringens var. egens Pr^vot, Ann. Inst. Past., 61, 1938, 78). Clostridium perfringens Type B, Wils- don. (Bacillus of lamb dysentery, Dall- ing. Jour. Path, and Bact., 28, 1925, 536, and ibid., 29, 1926, 316; L. D. Bacillus, Dalling, Handbook Ann. Congr. Nat. Vet. Med. Assoc, Gt. Britain and Ire- land, 1928, 56; Bacillus welchii Type B, Wilsdon, Univ. Cambridge, Inst. Animal Path., 2nd Rept. of Dir., 1931, 73; Clos- tridium welchii (Type agni) Glenny et al., Jour. Path, and Bact., 37, 1933, 53; Ba- cillus agni Weinberg et al., Les Mic. Anaer., 1937, 233; Welchia agni Pr^vot, Ann. Inst. Past., 61, 1938, 78.) Clostridium, perfringens Type C, Wils- don. (Bacillus paludis McEwen, Jour. Compar. Path, and Ther., 43, 1930, 1; Bacillus welchii Type C, Wilsdon, Univ. Cambridge, Inst. Animal Path., 2nd Rept. of Dir., 1931, 73; Welchia agni var. paludis Pr^vot, Ann. Inst. Past., 61, 1938, 78; Welchia paludis Pr(5vot, Man. d. Class., etc., 1940,217.) Clostridium perfringens Type D, Wils- don. {Bacillus v)elchii Type D, Wilsdon, Univ. Cambridge, Inst. Animal Path., 2nd Rept. of Dir., 1931, 74; Bacillus ovitoxicus Bennetts, Austral. Inst. Sci. and Indus., Bull. No. 57, 1932, 5, and Vet. Jour., 88, 1932, 250; Welchia agni var. ovitoxicus Prevot, Ann. Inst. Past., 61, 1938, 78; Clostridium ovitoxicus Spray, in Manual, 5th ed., 1939, 773; Welchia ovi- toxicus Prevot, Man. d. Class., etc., 1940, 217.) Probably related (or possibly identical) varieties: Bacille du rhumatisme, FAMILY BACILLACEAE 791 Achalme, Compt. rend. Soc. Biol., Paris, 4S, 1891, 651, and Ann. Inst. Past., 11, 1897, 848 (Bacille and bacterium d'Achalme, Thiroloix, Compt. rend. Soc. Biol., Paris, 49, 1897, 268; Bacillus achal- mei Neveu-Lemaire, Prdcis Parasitol. Hum., 5th ed., 1921, 24); Bacillus em- physematis maligni Wicklein, Arch. f. Path. Anat. u. Physiol., 125, 1891, 91; Bacillus cadaveris Sternberg, Researches relating to the etiology and prevention of yellow fever, Washington, 1891, 212 {Bacterium cadaveris Migula, Syst. d. Bakt., 2, 1900, 510; not Bacillus cadaveris Klein, Cent. f. Bakt., I Abt., 23, 1899, 279; not Clostridium cadaveris Sternberg, loc. cit., 213; not Bacillus cadaveris Migula, loc. cit., 646); Bacillus cadaveris hutyricus Budaj', Cent. f. Bakt., I Abt., 24, 1898, 374 {Bacillus budayi and Bac- terium cadaveris butyricum LeBlaye and Guggenheim, Man. Prat, de Diag. Bact., 1914, 378; Eubacterium cadaveris Prevot, Ann. Inst. Past., 60, 1938, 295). Bacillus zoodysenteriae Weinberg et al., Les Mic. Anaer., 1937, 256 {Bacillus zoodysenteriae hungaricus Detre, Cent, f . Bakt., I Abt., Orig., 104, 1927, 251; Welchia perfringens var. zoodysenteriae Prevot, Ann. Inst. Past., 61, 1938, 78). Clostridium perfringens var. anaero- genes Hauduroy et al.. Diet. d. Bact. Path., 1937, 122 (Unnamed species of Grootten, Compt. rend. Soc. Biol., Paris, 100, 1929,499). Rods: Short, thick, 1.0 to 1.5 by 4.0 to 8.0 microns, occurring singly and in pairs, less frequently in short chains. Non- motile. Spores oval, central to excentric, not swelling rods. Encapsulated. Gram- positive. Gelatin: Liquefied and blackened. Agar surface colonies (anaerobic ) : Circular, moist, slightly raised, opaque center, entire. Broth: Turbid; peptolytic. Clearing with viscid sediment. Litmus milk: Acid, coagulated. Clot torn with profuse gas formation, but not digested. Potato: Thin, grayish-white streak; gas in subtended liquid. Indole not formed. Nitrites produced from nitrates. Acid and gas from glucose, fructose, galactose, mannose, maltose, lactose, sucrose, xylose, trehalose, raffinose, starch, glycogen and inositol. Mannitol not fermented. Salicin rarely fer- mented. Action on inulin and glycerol variable. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Egg-meat : Profuse gas production in 8 hours. The meat is reddened and the liquid becomes turbid. No digestion. Pathogenic for guinea pig, pigeon and mouse. Produces an exotoxin for which an antitoxin can be prepared. Optimum temperature 35°C to 37°C. Can grow at 50°C. Anaerobic. Distinctive characters : Stormy fer- mentation of milk, combined with non- motility. Source: Gaseous gangrene, feces, milk and soil. Habitat: Widely distributed in feces, sewage and soil . 25. Clostridium sphenoides (Bulloch etal.) Bergey etal. {Bacillus sphenoides Bulloch, Bulloch, Douglas, Henry, Mcintosh, O'Brien, Robertson and Wolf, Med. Res. Counc, Spec. Rept. Ser. No. 39, 1919, 43; Douglasillus sphenoides Heller, Jour. Bact., 7, 1922, 5; Bergey et al., Manual, 1st ed., 1923, 331; Plectri- dium sphenoides Prevot, Ann. Inst. Past., 61, 1938, 88.) From Greek, wedge- shaped. Described from Bulloch et al., loc. cit., as amplified by Hall, Jour. Inf. Dis., 30,1922,502. Rods: Small, fusiform in vegetative state, occurring singly, in pairs and occa- sionally in short chains. Sporulating cells cuneate. Motile. Spores spheri- cal, subterminal, becoming terminal on 792 MANUAL OF DETERMINATIVE BACTERIOLOGY maturation, swelling rods. Gram-posi- tive only in young cultures. Gelatin : Not liquefied. Agar surface colonies (anaerobic ) : Circular, or slightly irregular, entire. Blood agar surface colonies (anaerobic) : Minute dew-drops, becoming whitish, opaque. Blood is hemolyzed. Deep agar colonies : Minute, opaque, smooth disks. Broth: Turbid. Litmus milk: Acid; slowly and softly coagulated. Clot not digested. Indole not formed (indole formed by Tholby strain, Stanley and Spray, Jour. Bact., 4^, 1941, 256). Nitrites produced from nitrates. Acid and gas from glucose, galactose, maltose, lactose and salicin. Inulin, glycerol and dulcitol not fermented. Strains are apparently variable on manni- tol, sucrose, dextrin and starch. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Non-pathogenic for guinea pig and rabbit. Optimum temperature not determined. Grows well at 30°C to 37°C. Anaerobic . Source: From gangrenous war wounds. Habitat : Not determined, other than this source. 26. Clostridium innominatum Prevot. (Bacillus E, Adamson, Jour. Path, and Bact., 22, 1918-19, 391; Pr(5vot, Ann. Inst. Past., 6i, 1938,85.) From Latin, remaining unnamed. Rods: Very small, thick, tapering at one or both ends, occurring singly, paired, in chains and filaments. Involution forms abundant on glucose agar. Motile. Spores small, spherical, subterminal, swelling rods. Gram-positive, quickly becoming Gram-negative. Gelatin : Not liquefied. Glucose agar surface colonies (anaero- bic) : Two forms are produced : 1) Circu- lar, entire edge, opaque; 2) Diffuse, spreading, irregular and translucent. Plain agar surface colonies (anaerobic) : Small, circular, entire edge, whitish- translucent, becoming opaque-yellowish with age. Plain broth : Moderate turbidity, clear- ing by sedimentation in 3 to 4 days. Glucose broth : More abundant tur- bidity and slight gas production. Milk: Slowly acidified but not clotted. No further change. Glucose, maltose, lactose and mannitol fermented with acid and gas. Sucrose not fermented. Coagulated albumin : Not digested or blackened. Meat medium : Not digested or black- ened. Blood serum : Not digested or black- ened. Brain medium : Not digested or black- ened. Non -pathogenic (Prevot, loc. cil.). Grows well at 37°C. Anaerobic. Source : From septic and gangrenous war wounds. Habitat : Not determined, other than this source. 27. Clostridiiun filiforme Bergey et al. (Bacillus regularis filiformis Debono, Cent. f. Bakt., I Abt., Grig., 62, 1912, 234; Bergey et al., Manual, 1st ed., 1923, 331.) From Latin, thread-like. Rods : 0.5 to 0.8 by 3.0 to 5.0 microns, slender, occurring singly, in pairs, in chains and filaments. Non-motile. Spores very small, spherical, subterminal, or occasionally terminal, not swelling rods. Gram -positive. Gelatin : Not liquefied. Deep gelatin colonies: Small, gray, filamentous. Deep agar colonies : Irregular, gray, translucent, filamentous. Broth: Uniform turbidity. Litmus milk: Acid, but no further change. FAMILY BACILLACEAE 793 Potato : Gray, filamentous ; substance not digested. Acid and gas from glucose and lactose. Acid only from sucrose and dulcitol. Starch not fermented. Coagulated albumin not liquefied. Grows in gelatin at 22°C. Anaerobic. Source : From human feces. Habitat: Not determined, other than this source. 28. Clostridium sartagoformiim Partan- sky and Henry. (Jour. Bact., 30, 1935, 570.) From Latin, shaped like a fry- ing-pan. Rods : 0.3 to 0.5 by 3.5 to 6.0 microns. Slender, curved, with rounded ends, oc- curring singly. Motile. Spores oval, terminal, swelling rods. Gram-positive. Gelatin : Not liquefied. Agar surface colonies (anaerobic ) : Convex, discrete, circular, transparent to white and opaque. Surface moist and smooth. Blood agar not hemolyzed. Deep agar colonies : Regular, lenticular, smooth. Broth: Clear; no growth. Glucose broth: Turbid, with gas bub- bles. Litmus milk: Acid; slowly coagulated, with some gas formation. Clot not di- gested. Potato : Very scant growth. No gas in surrounding liquid. Indole not formed. Nitrites not produced from nitrates. Acid and gas from xylose, glucose, fruc- tose, galactose, sucrose, lactose, maltose, raffinose, inulin, salicin, mannitol, ace- tate and butyrate. Starch, ethanol, glyc- erol and dulcitol not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Scant growth. Brain medium not blackened or di- gested. Some gas is formed. Optimum temperature 37°C. Anaerobic . Distinctive character : Ferments sulfite waste liquor in 40 per cent concentration, forming butyric and acetic acids, H2 and CO2. Source : From garden soil and from stream and lake mud. Habitat: Not recorded; obviously soil. Distribution undetermined. 29. Clostridium paraputrificum (Bien- stock) Snyder. (Art V, Bienstock, Fortschr. d. Med., 1, 1883, 612; Bacillus diapthirus Trevisan, I generi e le specie delle Batteriacee, 1889, 15; Bacillus para- putrificus Bienstock, Ann. Inst. Past., 20, 1906, 413, and Strassburger Med. Zeit., 3, 1906, 111; Bacillus paraputrificus coli Henry, Brit. Med. Jour., 1, 1917, 763; Tissierillus paraputrificus Heller, Jour. Bact., 7, 1922, 27; Snyder, Jour. Bact., 32, 1936, 401 ; Plectridium paraputrificum Prevot, Man. d. Class., etc., 1940, 160.) From Latin, also putrefying. Probable synonyms : Kopfchenbak- terien, Escherich, Fortschr. d. Med., 8, 1885, 515; Bacillus No. 3, Rodella, Ztschr. f. Hyg., 39, 1902, 209 {Plectriclium fluxum Prevot, Ann. Inst. Past., 61, 1938, 87); Art XI, Hibler, Cent. f. Bakt., I Abt., 25, 1899,516 (Art IX, Hibler, Untersuch. ii. d. Path. Anaer., 1908, 3 and 407; Plectridium nonum Prdvot, Ann. Inst. Past., 61, 1938, 88) ; Anaerobe b, Dalyell, Jour. Path, and Bact., 19, 1914-15, 281; Bacillus innutritus Kleinschmidt, Mo- natschr. f. Kinderheilkunde, 62, 1934, 18 {Palmula innutrita Prevot, Ann. Inst. Past., 61, 1938, 89; Acufonuis innutritus Prevot, Man. d. Class., etc., 1940, 165). Described from Hall and Snyder, Jour. Bact., 28, 1934, 181. Rods : 0.3 to 0.5 by 2.0 to 6.0 microns. Straight or slightly curved, single, in pairs, or in short chains. Ends rounded. Motile with peritrichous flagella. Spores oval, terminal, swelling rods. Gram- positive. Gelatin : Not liquefied. Gas is formed. Blood agar surface colonies (anaerobic ) : Delicate, irregular, round-topped dew- drops. Non-hemolytic. 794 MANtTAL or DETERMINATIVE BACTERIOLOGY Deep agar colonies: Small, irregular, opaque, dense, cottony masses. Gas is formed. Broth : Diffuse turbidity. Milk: Usually coagulated in from 6 to 10 days. Abundant gas, but no peptoni- zation. Indole is not formed. Acid and gas from glucose, fructose, galactose, maltose, lactose, sucrose, raffi- nose, dextrin, soluble starch, amygdalin and salicin. Xylose, inulin, mannitol and glycerol not fermented. Coagulated albumin not liquefied. Blood serum not liquefied or discolored. Brain medium not blackened or di- gested. Non-proteolytic. Non-pathogenic for guinea pig and rabbit. Grows well at 37°C. Anaerobic. Source: Feces, gaseous gangrene, and postmortem fluid and tissue cultures. Habitat: Undetermined, other than these sources. Evidently occurs com- monly in intestinal canal of human beings. 30. Clostridium cochlearium (Bulloch etal.) Bergeyetal. (Bacillus Type IIIc, Mcintosh, Med. Res. Counc, Spec. Rept. Ser. No. 12, 1917, 20; Bacillus cochlcarius Bulloch, Bulloch, Douglas, Henry, Mcin- tosh, O'Brien, Robertson and Wolf, Med. Res. Ck)unc., Spec. Rept. Ser. No. 39, 1919, 40; Flemingillus cochlcarius HeWer, Jour. Bad., 7, 1922, 5; Bergey et al.. Manual, 1st ed., 1923, 333; Plectridium cochlear- ium Pr^vot, Ann. Inst. Past., 61, 1938, 88; Plectridium incertum. Prevot, idem.) From Latin, spoon-shaped. Rods: Slender, straight, occurring chiefly singly , or infrequently in pairs and in short chains. Motile with peritrichous flagella. Spores oval, terminal, swelling rods. Weakly Gram-positive. Gelatin : Not liquefied. Agar surface colonies (anaerobic ) : Circular, clear, entire, or with crenated edge. Deep agar colonies : Lenticular, entire. Broth: Turbid. Litmus milk: Unchanged. Glucose not fermented. Carbohydrates not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Meat medium : Slightly reddened. Not blackened or digested. Little gas of non- putrefactive odor. Non-pathogenic. Optimum temperature 30°C to 35°C. Anaerobic. Source : From human war wounds and septic infections. Habitat: Not determined, other than these sources. Probably occurs in soil. 31. Clostridium kluyveri Barker and Taha. (Jour. Bact., IfS, 1942, 347.) Named for A. J. Kluyver, in whose labora- tory the organism was discovered. Rods : 0.9 to 1.1 by 3.0 to 11.0 microns. Straight to slightly curved; usually sin- gle, but also paired and occasionally in long chains. Motile with peritrichous flagella. Spores oval, terminal, swelling rods. Generally Gram-negative; some strains weakly Gram -positive when young. Iron-gelatin (Spray) : No growth. Surface agar colonies (anaerobic ) : Growth slow and restricted by residual traces of oxygen. Rough and smooth colonies are produced. Deep agar colonies (j^east autolysate and C2H5OH): Small colonies (1 to 3 mm) after 2 to 3 days; two types are formed : a) fluffy spheres with dense nuclear center and filamentous periphery ; b) compact, lenticular colonies. Little gas is formed. Plain broth: No growth. Glucose broth : No growth. Milk or iron-milk (Spray) : No growth. Indole not recorded (probably nega- tive). Nitrites not recorded (probably nega- tive). Glucose not fermented. Carbohydrates not fermented. FAMILY BACILLACEAE 795 Cellulose not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not digested or black- ened. Probably non-pathogenic. Optimum temperature about 34°C. Grows between 19°C and 37°C. Optimum reaction about pH 6.8. Range for growth pH 6.0 to 7.5. Anaerobic . Distinctive characters : Large size of cells, and slow growth, accompanied by non-putrefactive odor of caproic acid and of higher alcohols. Growth is exception- ally favored by synergistic association with Methanobacteriutn omelianskii. In pure culture a high concentration of yeast autolysate is required. Caproic acid is formed from ethyl alcohol . Source : From black mud of fresh water and marine origin. Habitat: Not determined, other than these sources. Evidently widely dis- persed in nature. 32. Clostridium acidiurici (Liebert) Barker. {Bacillus acidi urici Liebert, Koninkl. Akad. v. Wetensch., Proc. Sect. Sci., Amsterdam, 12, 1909, 55; Barker, Jour. Bact., 36, 1938, 323.) Named from its characteristic ability to ferment uric acid. Rods : 0.5 to 0.7 by 2.5 to 4.0 microns; straight. Motile with peritrichous fla- gella. Spores oval, terminal, swelling rods. Most strains Gram-negative. A few strains weakly Gram-positive, quickly becoming Gram-negative. Iron-gelatin (Spray) : No growth. Deep plain agar : No growth. Deep uric acid agar colonies: Whitish, compact, lobate, 1 to 2 mm in diameter, with irregular edge; surrounded by a temporary zone of precipitated am- monium ureate which gradually disap- pears. Surface uric acid agar colonies (anaero- bic) : Variable with strain and with moisture of medium. Colonies 1 to 2 mm in diameter, opaque, white, raised. round, smooth edge, with concentric sur- face markings, and of rubbery consist- ency. Other colonies may be very thin, soft, transparent, with fimbriate projec- tions, spreading to cover almost the entire plate. Intermediate colony types also observed. Plain broth : No growth. Glucose broth : No growth. Iron-milk (Spray) : No growth. Indole not recorded (probably nega- tive). Nitrites not recorded (probably nega- tive). Glucose not fermented. Carbohydrates not fermented. Cellulose not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not digested or black- ened. Probably non-pathogenic. Optimum temperature about 35°C. Optimum reaction about pH 7.5; lower limit for growth about pH 6.5. Anaerobic. Distinctive characters : Requires uric acid, or certain other purines, as a primary source of carbon and energy. The purines are converted mainly into ammonia, CO2 and acetic acid. During growth the medium tends to become alka- line (pH 8.0 to 8.5); there is no visible evolution of gas. Source: From soils of diverse origin. Habitat : Evidently widely dispersed in soils. Present in fecal material of yellow-shafted flicker (Colaptes auratus). 33. Clostridium capitovale (Snyder and Hall ) Snyder. (Bacillus capitovalis Sny- der and Hall, Cent. f. Bakt., I Abt., Orig., 135, 1935, 290; Clostridium capi- tovalis Snyder, Jour. Bact., 32, 1936, 401; Plectridium capitovalis Pr^vot, Ann. Inst. Past., 61, 1938, 87.) From Latin, oval -headed. Rods : 0.5 to 0.8 by 2.0 to 2.5 microns. Slender, commonly curved, with rounded ends, occurring singly, in pairs, and rarely in short chains. Motile with long peri- 796 MANUAL OF DETERMINATIVE BACTERIOLOGY trichous flagella. Spores oval, terminal, swelling rods. Gram-positive. Gelatin : Liquefied. Blood agar surface colonies (anaerobic) : Tiny, transparent, round or irregular dew-drops, becoming opaque. Non-he- molytic. Deep agar colonies: Small, opaque, lenticular to heart-shaped. Tryptone broth : Turbid. Gas is formed . Milk: Often, but not invariably, clotted. Acid is formed. Clot, when formed, is not digested. Indole not formed. Nitrites not produced from nitrates. Acid and gas from glucose, fructose and galactose. Maltose, lactose, sucrose, raf- finose, xylose, inulin, dextrin, starch, cellulose, amygdalin, salicin, mannitol and glycerol not fermented. Coagulated albumin liquefied. Blood serum slowly softened and par- tially liquefied. Not blackened. Mildly proteolytic. Brain medium is blackened; slightly softened, but not conspicuously liquefied. Pathogenicity : Guinea pig may show slight subcutaneous edema; usually no effect. Non-pathogenic for rabbit. Grows at 37°C. Anaerobic. Source : Human feces, gaseous gangrene and septicemia. Habitat: Not determined, other than these sources. 34. Clostridium parabifermentans comb. nov. {Bacillus jparahifermentans sporogenes de Gasperi, Compt. rend. Soc. Biol., Paris, 67, 1909, 492.) From Greek, para, near, and Latin, bis, twice, and fermentum, a ferment. Rods : 0.5 to 0.7 by 4.0 to 5.0 microns, occurring singly, in pairs and in chains of 3 to 5 cells. Motile. Spores oval, ter- minal, swelling rods. Gram-positive. Glucose gelatin : Rapid growth with liquefaction. Deep glucose agar colonies : Lenticular, regular, opaque, whitish. Agar dis- rupt ed by considerable gas of putrefactive odor. Glucose broth : Abundant growth with uniform turbidity and with viscous sedi- ment. Milk: Acidified but not coagulated. Casein slowly precipitated with slow, but complete, digestion. Indole formed in trace. Glucose, lactose and sucrose fermented to acids. (Gas not recorded.) Starch is not fermented. Coagulated albumin actively liquefied. Non-pathogenic for mouse. Grows between 22°C and 37°C. Anaerobic. Source : From putrefying game (pheas- ant and guinea-fowl). Habitat : Undetermined, other than this source. 35. Clostridixim ovalare Bergey et al. {Bacillus putrificus ovalaris Debono, Cent. f. Bakt., I Abt., Grig., 62, 1912, 231; Bergey et al., Manual, 1st ed., 1923, 336; Plectridium ovalaris Pr^vot, Ann. Inst. Past., 61, 1938, 88.) From Latin, oval. Rods : 0.3 to 0.4 by 6.0 to 8.0 microns, straight or curving, ends rounded, occur- ring singly, in pairs and in short chains. Motile. Spores oval, terminal, swelling rods. Gram-positive. Gelatin : Rapidly liquefied. Deep glucose agar colonies : Small, globular, entire, becoming brownish. Scant gas is formed. Broth: Turbid. Litmus milk : Acid, peptonized without coagulation. Indole not formed. Acid and scant gas from glucose and lactose. Acidonly from sucrose. Dulci- tol not fermented. Coagulated albumin rendered transpar- ent, then slowly peptonized, with a putrefactive odor. Grows at 22°C and at 37°C. Anaerobic . Source: Originally from putrid meat, later from feces. FAMILY BACILLACEAE 797 Habitat : Not determined, other than these sources. 36. Clostridiiun zoogleicum Bergey et al. {Bacillus sporogenes zoogleiciis Dis- taso, Cent. f. Bakt., I Abt., Orig., 59, 1911, 99; Bergey et al.. Manual, 1st ed., 1923, 335.) From Greek, zoogleal. Rods : Fairly long, occurring singly, in pairs and in short chains. Motile. Spores large, oval, terminal, swelling rods. Gram-positive. Gelatin : Growth and liquefaction not recorded. Deep agar colonies: Small, gray, slightly opaque, becoming heart-shaped. Gas is not formed. Broth: Turbid, then clearing with zoogleal sediment. Litmus milk: Slowly coagulated, then digested. Litmus reduced. Indole is formed in trace. Acid but no gas from glucose. Lactose and sucrose not fermented. Coagulated albumin liquefied, leaving a clear fluid and zoogleal sediment . Grows at 37°C. Anaerobic. Source : From human feces. Habitat : L'ndetermined, other than this source. 37. Clostridium thermosaccharolyticum McClung. (Jour. Bact., 29, 1935, 200; Terminosporus thernwsaccharolyticus Prevot, Ann. Inst. Past, 61, 1938, 86.) From Greek, heat, and sugar-digesting. Rods: 0.4 to 0.7 by 3.5 to 7.5 microns, slender, granulated, occurring singly and in pairs, not in chains. Motile with peri- trichous flagella. Spores spherical, ter- minal, swelling rods. Gram -negative. Gelatin : Not liquefied. Pea-infusion agar surface colonies (anaerobic): Granular, grayish-white, raised center, with feathery edges. Deep glucose-tryptone agar colonies : Small, lenticular, smooth. Liver-infusion broth over liver meat : Turbidity and gas. Litmus milk: Litmus reduced. Acid and slow but firm coagulation; coagulum split with gas. Clot not digested. Indole not formed. Nitrites not produced from nitrates. Cellulose not fermented. Acid and gas from arabinose, fructose, galactose, glucose, mannose, xylose, cello- biose, lactose, maltose, sucrose, trehalose, dextrin, glycogen, corn-starch, amygda- lin, esculin, a-methyl glucoside and salicin. Raffinose weakly fermented. Rhamnose, inulin, pectin, erythritol, inositol, mannitol, glycerol, quercitol and Ca-lactate not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Meat-medium not blackened or di- gested. Non-pathogenic on feeding to white rat, or by injection into rabbit. Optimum temperature 55°C to 62°C. Thermophilic. Anaerobic. Source : From hard-swell of canned goods, and from soil. Habitat: Not determined, other than these sources. 38. Clostridium caloritolerans Meyer and Lang. (Jour. Inf. Bis., 39, 1926, 321 ; Plectridium caloritolerans Prevot, Ann. Inst. Past., 61, 1938, 87.) From Latin, heat-enduring. Rods : 0.5 to 0.8 by 8.0 to 10.0 microns, with rounded ends, occurring singly, in pairs, in chains and in curved filaments. Motile with peritrichous flagella. Spores spherical or pear-shaped, terminal, swell- ing rods. Gram-positive. Gelatin : Not liquefied. Glucose blood agar surface colonies (anaerobic): Small, flat, grayish, rhizoi- dal. Non-hemolytic. Deep liver agar colonies: Small, flat, transparent disks with large polar tufts. Some colonies become flufi^y. Broth: Slight turbidity. Glucose broth : Abundant turbidity. 798 MANUAL OF DETERMINATIVE BACTERIOLOGY with clearing by sedimentation. Gas is formed. Brom cresol purple milk : No change. Indole not formed. Acid and gas from glucose, galactose and maltose. Fructose feebly fer- mented. Lactose, sucrose, raffinose, inu- lin, salicin, mannitol, inositol and glyc- erol not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Beef -heart mash medium : Reddened ; not blackened or digested. Non-pathogenic for mouse, guinea pig and rabbit. Optimum temperature not determined. Grows at 37°C. Anaerobic . Source: From an old culture of Clos- tridium parabotulinum Type A. Habitat: Not determined, other than this single isolation. 39. Clostridium tetanoides (Adamson) Hauduroy et al. (Unnamed anaerobe, Adamson and Cutler, Lancet, 1, 1917,688; Bacillus tetanoides (A) Adamson, Jour. Path, and Bact., 22, 1918-19, 382; Hau- duroy et al., Diet. d. Bact. Path., 1937, 140; Palmula macrospora Pr^vot, Ann. Inst. Past., 61, 1938, 88; Acuformis ma- crosporus Prevot, Man. d. Class., etc., 1940, 166.) From Latin, tetanus-like. Rods : 1.0 to 2.0 by 4.0 to 12.0 microns (averaging 1.0 to 1.5 by 6.0 to 7.0 mi- crons), with rounded to slightly tapered ends, occurring singly, in pairs and in chains of 3 to 5 cells, but not in filaments. Motile only in young cultures. Spores large, spherical, terminal, swelling rods. Gram-positive in young cultures, soon becoming Gram-negative. Gelatin : Not liquefied. Glucose agar surface colonies (anaero- bic) : Circular, regular, opaciue, bluish- gray, moist, shining, thick, raised. Surface flat, Ijecoming conical in center with age. On moist medium showing radiating, dendritic branching. Growth becomes tenacious -mucoid. Plain agar surface colonies (anaerobic ) : Confluent, becoming an opaque film. Isolated colonies circular to slightly ir- regular. Dendritic branching and mu- coid tendency less evident than on glucose agar. Glucose agar stab : Thick growth along stab, starting 0.5 cm below surface. No gas or splitting of medium. Neutral-red glucose agar: Reduced to orange by transmitted, and greenish- fluorescent by reflected light. Plain broth: Early slight turbidity, with clearing and mucoid sedimentation. Glucose broth : Abundant turbidity and profuse mucoid sediment. Milk : Slight and slowly increasing alka- linity, with slow separation of casein. No further change. Indole : Trace formed in broth. Glucose and maltose fermented with acid but no gas. Lactose, sucrose, manni- tol, starch and cellulose not fermented. Coagulated albumin : Not digested or blackened. Meat medium : Not digested or black- ened. Blood serum : Not digested or black- ened. Brain medium : Not digested or black- ened. Non-pathogenic for guinea pig and rabbit. Optimum temperature not recorded. Grows well at 37°C. Anaerobic . Source: From war wounds, from post- mortem blood culture, and from garden soil. Habitat: Not determined, other than these sources. 40. Clostridium tetani (Fliigge) Hol- land. (Tetanusbacillen and Tetanuser- reger, Nicolaier, Deuts. Med. Wchnschr.. 10, 1884, 843; Bacillus tetani Fliigge, Die Mikroorg., 2 Aufl., 1886, 274; Pacinic nicolaieri Trevisan, I generi e le speci* delle Batteriacee, 1889, 23; Plectridium FAMILY BACILLACEAE 799 teLani Fischer, Jalirb. 1'. Wissensch. Botanik, S7, 1895, (147?); Holland, Jour. Bact., 5, 1920, 220; Nicollaierillus tetani Heller, Jour. Bact., 7, 1922, 7.) From tetanus, lockjaw. Rods : 0.4 to 0.6 by 4.0 to 8.0 microns, rounded ends, occurring singly, in pairs, and often in long chains and filaments. Motile with peritrichous flagella. Spores spherical, terminal, swelling rods. Gram-positive. Gelatin: Slowly liquefied and black- ened. Serum agar surface colonies (anaero- bic) : Small, transparent, villous to fim- briate margin. Blood agar is hemolyzed. Deep agar colonies : Fluffy, cottony spheres, usually without visible central nucleus. Broth : Slightly turbid. Gas is formed. Some strains clear quickly by sedimen- tation. Litmus milk: Slow precipitation of casein, or soft clotting. Clot slowly softened, but not definitely digested. Little gas is formed. Lidole is formed. Nitrites not produced from nitrates. Glucose not fermented. Carbohydrates not fermented. Coagulated albumin slowl}^ liquefied. Blood serum slowly softened, with feeble digestion. Brain medium blackened and slowly digested. Not actively proteolytic. Pathogenic and toxic. Forms a potent exotoxin for which an antitoxin is pre- pared. Toxin intensely toxic on injec- tion but not on feeding. Optimum temperature 37°C. Anaerobic. Source: Originally isolated from ani- mals inoculated with garden soil extract. Frequently isolated from wounds in hu- man tetanus. Habitat : Common in soils, and in hu- man and horse intestine and feces. 41. Clostridium lentoputrescens Hart- sell and Rettger. (Bacillus der Eiweiss- faulniss, Bienstock, Fortschr. d. Med., 1, 1883, 614 (Art IV, Bienstock, ibid., 612; Eiweissbacillus, Bienstock, Ztschr. f. klin. Med., 8, 1884, 38); Bacillus albu- minis Schroeter, in Cohn's Kryptogamen- Flora von Schlesien, 3, 1, 1886, 162; Bacillus putrificus coli Fliigge, Die Mikroorg., 2 Aufl., 1886, 303; Pacinia putrifica Trevisan, I generi e le specie delle Batteriacee, 1889, 23; Bacillus putrificus Bienstock, Ann. Inst. Past. 13, 1899, 861; Bacillus butyricus putre- faciens Rodella, Ann. Inst. Past., 19 1905, 804; Putribacillus vulgaris Orla- Jensen, Cent. f. Bakt., II Abt., £2, 1909, 343; Clostridium putrificum Holland, Jour. Bact., 5, 1920, 220; Putrificus bienstocki Heller, Jour. Bact., 7, 1922, 8; Bacillus putrificus (coli) Lehmannand Neumann, Bakt. Diag., 7 Aufl., 3, 1927, 661; Hartsell and Rettger, Jour. Bact., 27, 1934, 39 and 497; Plectridium putrifi- cum and Plectridium putrificum var. lentoputrescens Prevot, Ann. Inst. Past., 61, 1938, 88.) From Latin, slowly made putrid. Probable synonyms : Bacillus radiatus Liideritz, Ztschr. f. Hyg., 5, 1889, 149 {Cornilia radiata Trevisan, loc. cit., 22; Bacillus radiatus anaerobius Hopffe, Ztschr. f. Infkrnkh. u. Hyg. Haust., 14, 1913, 392); Bacillus cadaveris sporogenes {anaerobicus) Klein, Cent. f. Bakt., I Abt., 25, 1899, 279 {Bacillus cadaveris Klein, idem; not Bacillus cadaveris Sternberg, Researches relating to the etiology and prevention of yellow fever, Washington, 1891, 212; not Bacillus cadaveris Migula, Syst. d. Bakt., 2, 1900, 646; Bacillus cadaveris sporogenes Klein, loc. cit., 284; Plectridium cadaveris Pre- vot, Ann. Inst. Past., 61, 1938, 88); Art XIV, von Hibler, Untersuch. ii. d. Path. Anaer., 1908, 3 and 413; Bacillus tetan- oides (B) Adamson, Jour. Bact. and Path., 22, 1918-19, 388. Hartsell and Rettger, loc. cit., conclude that their organism differs very ma- terially either from Clostridium cochlear- ium or from Bacillus putrificus, as described by Cunningham, .Jour. Bact., 800 MANUAL OF DETERMINATIVE BACTERIOLOGY 24, 1932, 61, and, as it cannot be definitely related to any other anaerobic species (including the Eiweissbacillus, Bien- stock, loc. cit., Bacillus putrificus coli Fltigge, loc. cit., Bacillus putrificus Bien- stock, loc. cit., etc.), they propose the name of Clostridium, lentoputrescens for this species. Rods : 0.4 to 0.6 by 7.0 to 9.0 microns, with rounded ends, occurring singly, in pairs and in chains. Motile with peri- trichous flagella. Spores spherical, ter- minal, swelling rods. Weakly Gram- positive, becoming Gram -negative. Gelatin : Liquefied. Agar surface colonies (anaerobic) : Small, circular, flat, edge crenated to filamentous spreading. Develop a ground-glass appearance. Deep agar colonies : Fluffy spheres with fibrils radiating from a central nucleus. Blood agar is hemolyzed. Litmus milk: Slow, soft coagulation or flocculent precipitation. Casein is slowly digested. Indole is formed (Hall, Jour. Lif. Dis., 30, 1922, 141 ) . Not formed (Hartsell and Rettger, loc. cit., 509). Nitrites not produced from nitrates. Hydrogen sulfide formed in egg-meat medium. Carbohydrates not fermented. Glu- cose slightly attacked without distinct acid (Hartsell and Rettger, loc. cit., 508). Coagulated albumin slowlj' liquefied and blackened. Blood serum is liquefied. Gas is formed. Brain medium slowly blackened and digested. Egg-meat medium : Slightly turbid liquid. Meat reddened in 7 to 10 days, then digested with a foul odor. Non-pathogenic for white mouse, guinea pig and rabbit. Filtrate non-toxic on injection or feeding. Grows well at 37°C. Anaerobic. Source : From putrefying meat. Habitat: Intestinal canal of human. Widely dispersed in soil. 42. Clostridium filamentosum Bergey et al. {Bacillus putrificus filamentosus Distaso, Cent. f. Bakt., I Abt., Orig., 59, 1911, 98; Bergey et al.. Manual, 1st ed., 1923, 333; Palviula filamentosa Prevot, Ann. Inst. Past., 61, 1938, 88; Acuformis filamentosus Prevot, Man. d. Class., etc., 1940, 165.) From Latin, filamentous. Rods : Slender, occurring singlj^, in pairs and in chains. Motile. Spores spherical, or nearly so, terminal, swelling rods. Gram-positive. Gelatin : Liquefied. Deep glucose agar colonies : Delicate, cotton J' flocculi. Only a trace of gas formed . Broth : Turbid. Litmus milk : May or may not coagulate and digest slowly (variable). Indole formed in scarcely detectable trace. Odor of scatol. Glucose is feebly fermented, with little gas. Lactose and sucrose not fermented. Coagulated albumin : Rendered trans- parent, then slowly liquefied. Grows well at 37°C. Anaerobic . Source : From human feces. Habitat : Not determined, other than this source. 43. Clostridium tetanomorphum (Bul- loch et al.) Bergey et al. (Bacillus pseudo -tetanus, Type No. IX, — Tetanus- like Bacillus (Pseudotetanus Bacillus), Mcintosh and Fildes, Med. Res. Counc, Spec. Rept. Ser. No. 12, 1917, 11 and 32; Bacillus tetanomorphus Bulloch et al., Med. Res. Counc, Spec. Rept. Ser. No. 39, 1919, 41; Macintoshillus tetano- morphus Heller, Jour. Bact., 7, 1922, 5; Bergey et al.. Manual, 1st ed., 1923, 330; Plectridium tetanomorphum Prevot, Ann. Inst. Past., 61, 1938, 87.) From Greek, shaped like the tetanus organism. Synonyms or possibly related species : Bacillus pscudotetani Migula, Syst. d. Bakt., 2, 1900, 598 (Tetanusahnlicher Ba- cillus and Pseudotetanusbacillus, Tavel and Lanz, Mitteil. a. klin. Med. Inst. d. Schweiz, 1, 1893, 162; Bacillus taveli FAMILY BACILLACEAE 801 Chester, Man. Determ. Bact., 1901, 304; Plectridium pseudotetanicum Pr^vot, Ann. Inst. Past., 61, 1938, 87; Plec- tridium pseudo-tetanicurn Pr^vot, Man. d. Class., etc., 1940, 158) ; possibly identical with Bacillus fragilis Veillon and Zuber, Arch. d. M^d. Expt. et d'Anat. Path., 10, 1898, 536 and Bacillus ramosus Veil- lon and Zuber, ibid., 537. Rods: Slender, with rounded ends, oc- curring singly and in pairs, not in chains. Motile with peritrichous flagella. Spores spherical, or nearly so, terminal, swelling rods. Gram -positive. Gelatin : Not liquefied. Gelatin is liquefied (Hall, Jour. Inf. Dis., 30, 1922, 501). Agar surface colonies (anaerobic): Small, flat, irregularly circular, translu- cent, crenated. Deep agar colonies: Small, opaque, irregular; not woolly or branched. Agar slant (anaerobic) : Grayish, trans- lucent growth. Broth: Turbid. Litmus milk : Unchanged ; or occasional slight reduction of litmus. Acid and gas from glucose and maltose. Fructose, galactose, lactose, sucrose, sali- cin, inulin, mannitol and glycerol not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium not blackened or di- gested. Egg -meat medium : Slight gas formation in 48 hours. White crj^stals are depos- ited. Non-pathogenic for guinea pig and rabbit. Grows at 30°C and 37°C. Anaerobic . Source : Frqm war wounds and from soil. Habitat : Not determined other than these sources. Probably rather common in soil. 44. Clostridiixm alcaligenes Bergey et al. {Bacillus anaerobicus alcaligenes Debono, Cent. f. Bakt., I Abt., Orig., 62, 1912, 232; Bergey et al.. Manual, 1st ed., 1923, 331 ; Palmula alcaligenes Pr^vot, Ann. Inst. Past., 61, 1938, 89; Actiformis alcaligenes Pr^vot, Man. d. Class., etc., 1940, 165.) From French alcali, alkali and Latin suffi.x, producing. Rods : Long, slender, occurring singly, in pairs and in short chains. Non-motile. Spores spherical, terminal, swelling rods. Gram -positive. Gelatin: Not liquefied. Deep glucose agar colonies : Lenticular to irregular, or spherical, white, granular, entire. Broth : Uniform turbidity. Fecal odor. IMilk: Alkaline; casein slowly precipi- tated, with yellowish supernatant fluid. Indole is formed in abundance. Acid and gas from glucose and lactose. Sucrose and dulcitol not fermented. Cultures have odor of valerianic acid. Grows at 22°C and at 37°C. Anaerobic. Source : From human feces. Habitat : Not determined, other than this source. 45. Clostridium angulosum (Distaso) Hauduroy et al. (Bacillus angulosus Distaso, Cent. f. Bakt., I Abt., Orig., 62, 1912, 439; not Bacillus angulosus Garnier and Simon, Bull, et Mem. Soc. Mdd. Hop. Paris, 2^, 1907, 1034; Bacteroides angulo- sus Bergey et al., Manual, 1st ed., 1923, 260; Hauduroy et al.. Diet, des Bact. Path., 1937, 91.) From Latin angulosus, having angles, hooked. Rods : Short, thick, with rounded ends, occurring singly and in pairs. Long rods sometimes bent to form an obtuse angle. Encapsulated. Non -motile. Spores verj' small, spherical, terminal, slightly swelling rods. Gram stain not recorded. Plain gelatin : No growth at 20°C or at 37°C. Glucose gelatin : Grows well at 37°C. Growth cloudy at first, then clears and liquefies, with whitish, powdery precipi- tate. Glucose agar deep colonies : Large, 802 MANUAL OF DETERMINATIVE BACTERIOLOGY angular, opaque, yellowish. Gas bubbles are formed. Broth : Turbid. Litmus milk: Acid and coagulated in 14 days. Indole is formed. Acid and gas from glucose, lactose and sucrose. Butyric acid is formed. Coagulated albumin not liquefied. Odor of skatol. Optimum temperature 37°C. Anaerobic. Distinctive character : Resembles the Bacille neigeux, Jungano, Compt. rend. Soc. Biol., Paris, 62, 1907, 677, in form, but not in other respects. Source : From human feces. Habitat: Not determined, other than this source. 46. Clostridium putrefaciens (Mc- Bryde) Sturges and Drake. {Bacillus 'pulrejaciens McBryde, U. S. D. A., Bur. An. Ind., Bull. 132, 1911, 6; Sturges and Drake, Jour. Bact., U, 1927, 175; Pal- mula putrefaciens Pr^vot, Ann. Inst. Past., 61, 1938, 89 ; Acuformis putrefaciens Pr^vot, Man. d. Class., etc., 1940, 165.) From Latin, putrefying. Description from McBryde {loc. cit.) and amplified from Sturges and Drake {loc. cit.). Rods : 0.5 to 0.7 by 3.0 to 15.0 microns, rounded ends, occurring singly, in pairs, and in chains and filaments. Non-motile. Spores spherical, terminal, swelling rods. Gram-positive. Gelatin : Liquefied. Agar surface colonies (anaerobic ) : Small, filamentous. Agar slant (anaerobic) : Scanty, white, beaded, glistening growth. Broth: Moderate turbidity. Heavy, fiocculent sediment. Litmus milk: Rennet coagulation, peptonized. Litmus reduced. Indole not formed. Nitrites not produced from nitrates. Slight production of hydrogen sulfide. Acid and gas from glucose. Lactose, sucrose, maltose and starch not fer- mented. Coagulated albumin liquefied. Blood serum liquefied. Brain medium blackened and slowlj'' digested. Minced pork medium : Slight disinte- gration; sour, putrefactive odor. Non-pathogenic. Optimum temperature 20°C to 25°C. Slow growth at 0°C and no visible growth at 37°C. Anaerobic. Source : From muscle tissue of hogs at slaughter. Habitat: Not determined, other than this source. 47. Clostridium nigrificans Workman and Weaver. (Iowa State Coll. Jour. Sci., 2, 1927-28, 63; Werkman, Iowa State Coll. Research Bull. 117, 1929, 165.) From Latin niger, black and faciens, making. Rods : 0.3 to 0.5 by 3.0 to 6.0 microns, with rounded ends. Motile. Spores oval, subterminal, slightly swelling rods. Gram-positive. Gelatin : Not liquefied. Deep agar colonies : Show blackening of medium around colonies. Black in- creased by adding 0.1 per cent ferric chloride to medium. Milk: Not recorded. Indole not formed. Nitrites not produced from nitrates. Glucose not fermented. Carbohydrates not fermented. Coagulated albumin not liquefied. Blood serum not liquefied. Brain medium blackened but not digested. Hydrogen sulfide produced from cystine. Non-pathogenic to man, guinea pig, mouse, rat and rabbit. Optimum temperature 55°C. Ther- mophilic, growing at 65°C to 70°C. Anaerobic. Distinctive character : Black colonies in agar media. FAMILY BACILLACEAE 803 Source : From canned corn showing sulfur stinker spoilage; also occasionally from soil and manure. Habitat : Presumably soil, although de- tected with great difficulty. 48. Clostridium belfantii (Carboneand Venturelli) Spray. (Bacillus belfantii Carbone and Venturelli, Boll. 1st. Siero- ter., Milan, 4, 1925, 59; not Bacillus bel- /an though in young cultures very short, lightly curved rods may often pre- dominate. Size variable, even for the same strain, and strongly influenced by age of culture and composition of me- dium. Rather consistently short cells in young cultures in yeast extract, espe- cially when incubated anaerobically in the light, or in anaerobic cultures with substrates which permit only a slow" and scanty development, such as malonate. Dimensions in such cultures 0.6 to 0.8 bj- 1.2 to 2 microns. ^lore often, especially in older cultures, cells are much longer, up to 10 microns. Highly characteristic is the pronounced tendency to the forma- tion of irregularly shaped, bent and crooked long rods, occasionally swollen at one or both extremities, and fre- quently suggesting branching. Such cells usually form clusters reminiscent of Corynebactcrium and Mycobacteriutn cultures. Cells in young cultures actively motile by means of polar flagella; irregular and long cells as a rule non-motile. Gram- negative. Growth in liquid media never mucoid; sediment in older cultures homogeneous and smooth, readily redispersible. Color: Varies considerably, depending upon the medium, and especially in anaerobic illuminated cultures. Where development is slight (as in malonate, thiosulfate, and, usually, glycerol me- dia), the color is a light pink; in fatty acid-containing media more nearly dark reddish-brown. Color due to bacterio- chlorophyll and a number of different carotenoid pigments; most strains pro- duce in addition a water-soluble, non- carotenoid, bluish-red pigment which diffuses into the culture medium. In yeast extract cultures growth is possible over the range pH 6 to 8.5. With certain substrates, especially fatty acids, the combined effect of low pH and a substrate concentration of 0.1 to 0.2 per cent may prevent growth. No char- acteristic odors save that old cultures may develop a distinct ionone-likp frag- rance. Gelatin is not liquefied; leucine is generally utilized as a substrate. Most strains are able to grow on the surface of agar plates or slants; a few, especially when first isolated, appear more sensitive to oxygen and develop only in stabs in which the upper region may remain free of growth. Generally such strains can be adapted to grow at full atmospheric oxygen tension. Most fatty acids and hydroxj' acids are adequate oxidation substrates. All cultures can grow at the expense of thio- sulfate and produce rapid and profuse growth in glutarate and ethanol media. No development in media containing as the chief oxidation substrate 0.2 per cent sorbitol, glucose or mannose, even though these substances are not inhibi- tor^^ Molecular hj^drogen can be oxi- dized. All cultures can develop anaerobically in illuminated cultures bj^ photo- synthesis. Temperature optimum generally rather high, good development being possible up to 37 °C. However, certain strains exhibit a lower temperature optimum . Distinguishing characteristics: Mor- phological resemblance to species of Mycobacterium in old cultures; ability to grow with thiosulfate as the chief oxidizable substrate, and failure to develop in media which contain carbo- hydrates or sugar alcohols in a concen- tration of 0.2 per cent as the main oxidiz- able compounds. 864 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat: Regularly found in mud and stagnant bodies of water. Illustrations: Molisch, loc. cit., Plate I, fig. 1, 2; Plate II, fig. 10; van Niel, Loc. cit., fig. 1-3, p. 18, and fig. 18-26, p. 90. 2. Rhodopseudomonas gelatinosa (Mo- lisch) van Niel. {Rhodocystis gelatinosa Molisch, Die Purpurbakterien, Jena, 1907, 22; van Niel, Bact. Rev., 8, 1944, 98.) From Latin gclatio, freezing, indi- cating solidification, or in this case, clumping. Cells: In young cultures, short and small rods, approximately 0.5 by 1 to 2 microns. In old cultures much longer, up to 15 microns, and then irregularly curved rods, often swollen and gnarled in places up to 1 micron in width. In this stage the cells bear some resemb- lance to those found in old cultures of Rhodopseudomonas palustris, but the characteristic Mycobacterium-like clus- ters of the latter are absent. Single cells infrequent due to a copious mucus production in all media which causes the cells to clump together. While young cells are actively motile by means of polar flagella, motility is often diffi- cult to ascertain as a result of the pro- nounced tendency to conglomerate; the individuals in the clumps appear to be non-motile. Gram-negative. Gelatin is liquefied; of the single amino acids alanine, asparagine, aspartic and glu- tamic acids appear generally satisfactory substrates. Color: Quite distinctive in most an- aerobic cultures as a pale, delicate, pink- ish shade, rather peach-colored. Only in the presence of rather high concentra- tions of yeast extract (when a much heavier growth is obtained than with low concentrations supplemented with 0.2 per cent of various single oxidation sub- strates) do the slimy cell masses appear a dirty, faded brown. Color is due to bacteriochlorophyll and carotenoid pig- ments. Occasionally a water-soluble, non-carotenoid, bluish-red pigment is produced which diffuses into the culture medium. In yeast extract, growth occurs over a pH range extending from at least 6.0 to 8.5. Cultures produce a characteristic acrid odor. More sensitive to fatty acids than other species of Rhodopseudomonas; with 0.2 per cent propionate no growth occurs. The best single oxidizable substrates appear to be ethanol, glucose, fructose and mannose, as well as a variety of amino acids. Citrate also permits good growth; not, on the other hand, glycerol, mannitol, sorbitol or tartrate in the usual concentration of 0.2 per cent. Thiosulfate is not oxidized; behavior towards molecular hydrogen unknown. More pronouncedly microaerophilic than the other Rhodopseudomonas spe- cies; most cultures cannot develop on aerobically incubated slants or agar plates. Capable of strictly anaerobic develop- ment in illuminated cultures by virtue of a photosynthetic metabolism. Temperature relations so far unknown. Distinguishing properties: The small size of the individual cells, and the pro- nounced clumping which causes the cultures to be exceptionally stringy; the unusual color of the cell masses; the ability to liquefy gelatin, to utilize ci- trate and a number of amino acids. Correlated with these is the failure to grow in media with 0.2 per cent pro- pionate, tartrate and glycerol. Habitat: Regularly present in stagnant bodies of water and in mud. Illustrations: Molisch, loc. cit., Plate I, fig. 8; van Niel, loc. cit., fig. 55-60, p. 99; fig. 61-66, p. 100. 3. Rhodopseudomonas capsulatus (Mo- lisch) van Niel. {Rhodonostoc capsu- lalum Molisch, Die Purpurbakterien, Jena, 1907, 23; van Niel, Bact. Rev., 8, 1944, 92.) From Latin capsula, con- tainer (sheath). FAMILY ATHIORHODACEAE 865 Cells: Depending upon the pH of the medium, cells nearly spherical, or as dis- tinct rods, often devoid of motility. Motility due to polar flagella. The spherical cells are found in media with a pH below 7; they are usually arranged in chains resembling streptococci. Rod- shaped cells are characteristic for media with pH above 7; the higher the pH, the longer the rods. Individual cells slightly less than 1 micron wide, although attenuated rods (about 0.5 micron in width) are frequent at pH above 8, and slightly swollen cells (to 1.2 microns) are found in media containing sugars. Length varies from 1 to 6 microns; most common dimensions in approximately neutral media 2 to 2.5 microns. At pH above 8 abnormal growth in the form of irregular filaments. Outstandingly characteristic is the zigzag arrangement of the cells in chains. Cultures in media of pH 8 or above are distinctly mucoid. Gram-negative. Color: Anaerobic cultures develop with a brown color, the shade ranging from a light yellowish-brown to a deep mahogany brown. When grown in the presence of oxygen, the cultures are dark red. Even the pigmentation of the brown-colored organisms from an an- aerobic culture can be changed into a distinct red by shaking a suspension with air for some hours; light enhances the rate of this color change. Color due to bacteriochlorophyll and carotenoid pig- ments. Xo diffusible water-soluble pig- ment is produced. Growth possible over a pH range from at least 6 to 8.5, morphology becoming abnormal in the alkaline media. Most cultures are odorless, although occasionally a faint peach-like odor can be detected. Growth is not inhibited by the pres- ence of oxygen, although the pigmenta- tion is thereby affected. Fatty acids and most substituted acids are satisfactory substrates. Rapid and abundant growth with propionate at a concentration of 0.2 per cent. At this same concentration glutaric acid leads, at best, to very meager cultures, while tartrate, citrate and gluconate fail to induce growth, as do also ethanol, glycerol, mannitol and sorbitol. In media wdth 0.2 per cent glucose or fruc- tose good growth is obtained. No growth with mannose. Thiosulfate is not, but molecular hydrogen can be, oxi- dized by this species. Gelatin is not liquefied; of the amino acids alanine and glutamic acid are satis- factory substrates, while leucine is not utilized. Distinguishing properties: Cell shape and arrangement in chains; brown color of anaerobic, red pigmentation of aerobic cultures; abilitj^ to grovr in media with 0.2 per cent propionate, glucose, fruc- tose, alanine and glutamic acid; failure to develop with leucine, as well as with ethanol, glycerol, mannitol and sorbitol in the above-mentioned concentration. All cultures can develop anaerobically in illuminated cultures by a photosyn- thetic metabolism. Temperature optimum distinctly lower than for Rhodopseudomonas palustris, and, as a rule, around 25°C. Habitat: Regularly found in stagnant bodies of water and in mud. Illustrations: Molisch, loc. cit., Plate II, fig. 9; van Niel, loc. cit., fig. 4-6, p. 19; fig. 27-32, p. 92; and fig. 33-38, p. 93. 4. Rhodopseudomonas spheroides van Xiel. {Rhodococcus capsulatus Molisch, Die Purpurbakterien, Jena, 1907, 20 Rhcdococcus minor ^Molisch, ibid., 21 Rhodosphaera capsulata Buchanan, Jour Bact., 3, 1918, 472; Rhodosphaera minor Bergey et al., Manual, 1st ed., 1923, 405; Rhodorrhagus minor Bergey et al.. Manual, 3rd ed., 1930, 535; Rhodorrhagus capsulatus Bergey et al., Manual, 3rd ed., 1930, 535; van Xiel, Bact. Rev., 8, 1944, 95.) From Latin sphaera, a round body and Greek eidos, form of. 866 MANUAL OF DETERMINATIVE BACTERIOLOGY Cells: Generally single, nearly spheri- cal, diameter without slime capsule variable, depending upon medium, I'ang- ing from 0.7 to 4 microns. In young cultures actively motile by means of polar fiagella; motility soon ceases in media which are or become alkaline. Copious slime production in media at pH above 7. In strongly alkaline cul- tures abnormal cell-shapes occur in the form of irregular, swollen and distorted rods, often having the appearance of spore-bearing cells, simulated by the production of fat bodies. In sugar-con- taining media egg-shaped cells, meas- uring as a rule 2.0 to 2.5 by 2.5 to 3.5 microns, are frequently found. Gram- negative. Color : Anaerobic cultures develop with brown color, ranging in shade from a light, dirty greenish-brown to a dark brown. Cultures grown in the presence of oxygen are distinctly red. As in the case of Rhodopseudonionas capsulatus, the brown color of an anaerobic culture can be changed to red by shaking with air, light stimulating the color change. Color due to bacteriochlorophyll and carotenoid pigments. The large ma- jority of cultures of this species produces in addition a water-soluble, non-caro- tenoid, bluish-red pigment which diffuses into the culture medium. Gelatin is not liquefied, and growth with single amino acids appears some- what erratic. No definite correlations have been observed. Development is possible over a wide pH range, extending from at least 6.0 to 8.5. All cultures exhibit an unpleasant pu- trid odor. Requires for optimal development higher concentrations of yeast extract as a supply of growth factors than either Rhodopseudonionas palustris or Rhodo- pseudomonas capsulatus and is more sensitive to low fatty acid concentra- tions. With 0.2 per cent propionate in a neutral medium, no growth occurs; caproic and pelargonic acids are toxic in concentrations below 0.1 per cent. On the other hand, tartrate and gluconate can serve as oxidation substrates, as can also ethanol, glycerol, mannitol, sorbi- tol, glucose, fructose and mannose in 0.2 per cent concentrations. In sugar-containing media, acid is pro- duced; the pH may drop to below 4.0 before development ceases. Acid pro- duction from glucose occurs both in presence and absence of air, and in illuminated as well as in non-illuminated cultures. In cultures exposed to light, the acid usually disappears later on. Thiosulfate is not oxidized; hydrogen oxidation has not been observed. Oxygen does not prevent growth; colonies develop on the surface of agar plates exposed to air, with a red pig- mentation. Capable of strictly an- aerobic development in illuminated cul- tures by photosynthesis. Temperature optimum below 30°C. Distinguishing properties: Spherical cell-shape in most media; brown color of anaerobic and red pigmentation of aerobic cultures; growth with 0.2 per cent tartrate, gluconate, ethanol, glyc- erol, mannitol, sorbitol, glucose, fruc- tose and mannose; failure to grow with 0.2 per cent propionate. Habitat: Regularly found in stagnant bodies of water and in mud. Illustrations: Molisch, loc. cil., Plate II, fig. 15; van Niel, loc. cil., fig. 7-8, p. 19; fig. 39-45, p. 96; fig. 46-54, p. 97. Genus II. Rhodospirillum Molisch emend, van Nicl. (Molisch, Die Purpurbakterien, Jena, 1907, 24 ; van Niel, Bact. Rev., 8, 1944, 86; the genus now includes the genus Phaeospirillum Kluyver and van Niel, Cent. f. Bakt., II Abt., 9Jt, 1936, 396.) From Greek rhodon, red and M.L. spirillum, spirillum. Spiral-shaped bacteria, motile by means of polar fiagella. Gram-negative. FAMILY ATHIORHODACEAE 867 Contain bacteriochlorophyll and are potentially photosynthetic in the presence of extraneous oxidizable substances. Molecular oxygen is not produced. Unable to grow in strictly mineral media, even when possessed of the ability to utilize hydrogen as oxidizable substrate, due to the need for organic nutrilites. Produce accessory pigments causing the cultures, especially when grown in the light, to appear in various shades of red to brown. The type species is Rhodospirillum rubrum (Esmarch) Molisch. Key to the species of genus Rhodospirillum. I. Cultures red; cells well over 0.5 micron, usually about 1 to 1.2 microns in width. 1. Rhodospirillum rubrum. II. Cultures brown to orange; cells 0.5 micron or less in width. 2. Rhodospirillum fulvum. 1. Rhodospirillum rubrum (Esmarch) Molisch. [Spirillutn rubrum p]smarch. Cent. f. Bakt., 1, 1887, 225; Molisch, Die Purpurbakterien, Jena, 1907, 25; Rhodo- spirillum. photometricum Molisch, ibid., 24; Rhodospirillum giganteum Molisch, ibid., 24; Rhodospirillum longum Hama, Jour. Sci. Hiroshima Univ., Ser. B, Div. 2, 1, 1933, 135; Rhodospirillum gracilc Hama, ibid., 159.) From Latin ruber, red. Cells: Characteristically spiral- shaped, but size of elements variable within wide limits, depending upon en- vironmental conditions during growth. Width of cells from 0.5 to 1.5 microns; length from 2 to 50 microns, and over; even in a single culture such differences may be found. Also the shape and size of the spiral coil varies much; it usually ranges between 1 to 4 microns in width, and from 1.5 to 7 microns in length. In alanine media the majority of the cells occurs in the form of half-circles to com- plete rings; malate media tend to pro- duce much flattened spirals. In old cultures involution forms ap- pear, straightened spirals and irregularly swollen cells, the latter common in media with higher fatty acids. Such cells stain irregularly, contain fatty in- clusions, and are occasionally branched. Mucus is not produced. In calcium- delicient media the growth is flocculent, as if agglutinated. With an adequate calcium supply the growth in liquid media is homogeneous, suspended, and consists of single cells. Young cultures show active motility, due to polar flagella. Gram-negative. Gelatin is not liquefied; the amino acids alanine, asparagine, aspartic and glutamic acids are satisfactory oxidiz- able compounds. Color: Ordinarily deep and dark red, without any brownish tinge. In ethanol media lighter, and a characteristic pink. Pigment production markedly influenced by oxygen and light. Slants incubated in darkness present a pale grayish sur- face growth with a faint reddish hue, while often showing deep-red cell masses in the region between glass wall and agar surface where development proceeds at low oxygen tension. The color is due to bacteriochlorophyll and carotenoid pig- ments. Among the latter spirilloxan- thin is quantitatively predominant. Water-soluble, diffusible pigments are not produced. Development possible over a pH range of at least 6 to 8.5, although, as in other cases, the combination of an acid reac- tion and the presence of fatty acids may prevent growth. Cultures produce a distinctive odor, reminiscent of slightly putrid yeast. In general, grow well with fatty acids as the chief oxidizable substrate; how- ever, are prevented from growing by 0.2 per cent propionate in a neutral medium. Most substituted acids are equally satisfactory, with the exception 868 MANUAL OF DETERMINATIVE BACTERIOLOGY of tartrate, gluconate and citrate. In a concentration of 0.2 per cent, ethanol is a suitable substrate, whereas the carbo- hydrates and their corresponding poly- alcohols are not utilized. Thiosulfate is not oxidized; molecular hydrogen can be used by some strains. Rather microaerophilic ; many strains upon initial isolation incapable of growth at atmospheric oxygen tension. Subse- quent adaptation can be induced. But even such adapted cultures exhibit nega- tive chemotaxis to air. Capable of strictly anaerobic develop- ment in illuminated cultures on the basis of a photosynthetic metabolism. Temperature optimum generally be- tween 30° and 37°C. Distinguishing properties: The most important characteristics of the species are the spiral shape, combined with the ability to produce a red pigment with a definite absorption maximum at 550 millimicrons in the intact cells. Diag- nostically useful are the good growth in media with 0.2 per cent ethanol, alanine, asparagine, aspartate or glutamate, and the inadequacy of similar concentrations of carbohydrates and thiosulfate as substrates. Habitat: Regularly present in stagnant bodies of water and in mud. Illustrations: Molisch, loc. cil., Plate I, fig. 5-7; van Niel, Bact. Rev., 8, 1944, fig. 9-10, p. 19; fig. 11-16, p. 24; fig. 67- 75, p. 103; fig. 76-84, p. 104; fig. 85-90, p. 106; fig. 91-96, p. 107. 2. Rhodospirillum fulvum van Xiel. (Bact. Rev., 8, 1944, 108.) From Latin fulvurn., yellowish, tawny. Characteristic for the species is the very small size of the individual cells. These are not over 0.5 micron wide, and generally not longer than 2.5 microns. The most common shape consists of a complete turn of about 1 by 1.5 microns. In media with fatty acids as a substrate the spirals appear somewhat steeper than in fumarate, succinate or malate cul- tures. Swollen individuals resembling vibrios are encountered in cultures which do not appear quite healthy. Formation of mucus or clumping has not been ob- served. Gelatin is not liquefied; aspartate has been the only amino acid capable of in- ducing growth. Thiosulfate is not oxi- dized. Color: Quite distinct from that of Rhodospirillum rubrum; colonies and stab cultures are a reddish-brown, while liquid cultures often appear brownish- orange. The color is due to bacterio- chJorophyll and carotenoid pigments; among the latter spirilloxanthin, as evi- denced by the absence of an absorption maximum at 550 millimicrons, is not represented as a major constituent. Does not produce water-soluble, diffus- ible pigments. Capable of strictly anaerobic develop- ment in illuminated cultures, due to photosynthetic metabolism. Fatty acids and the four-carbon dicar- boxylic acids are uniformly good sub- strates; glutarate is not used. Ethanol and glucose, in a concentration of 0.2 per cent, have yielded satisfactory cultures; other carbohydrates, as well as the corre- sponding polyalcohols, have given nega- tive results. Little information available concern- ing pH and temperature relations. Be- haves generally as a strict anaerobe; adaptation to microaerophilic conditions has not been achieved. Negative aero- taxis very pronounced. Distinguishing properties: The small size and the color of the cultures serve as adeciuate criteria for its differentia- tion from Rhodospirilluui rubrum. The strictl}^ anaerobic nature and the failure to grow with glutarate and various amino acids except aspartate can prob- ably be used as supplementary specific properties. Habitat: Bodies of stagnant water and mud. Illustrations: Van Xiel, loc. cit., fig. 97-102, p. 109. FAMILY CHLOROBACTERIACEAE 869 FAIVIILY III. CHLOROBACTERIACEAE GEITLER AND PASCHER.* (Cyanochloridinae-Chlorobacteriaceae Geitler and Pascher, Die Siisswasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 451; Chlorothiobacteria Bavendamm, Ergeb. Biol., 13, 1936, 49.) Green bacteria, usually of small size, occurring singly or in cell masses of various shapes and sizes, developing in environments containing rather high concentrations of hydrogen sulfide and exposed to light. As a rule not containing sulfur globules but frequently depositing elementary sulfur outside the cells. Contain green pig- ments of a chlorophyllous nature, though not identical with the common green plant chlorophylls nor with bacteriochlorophyll. Capable of photosynthesis in the pres- ence of hydrogen sulfide; do not liberate oxygen. A number of genera have been proposed, characterized by special colonial growth forms, others on the basis of a supposed s3-mbiotic habitus, where the green bacteria grow in more or less characteristic aggregates together with other micro-organisms. In view of the variations in size and shape exhibited by the only member of this group which has so far been obtained and studied in pure culture (van Xiel, Arch. f. Mikrobiol., 3, 1931, 65fT.) the validity of many of these genera is doubtful. The fol- lowing keys and descriptions, therefore, bear a strictly provisional character. Here, as in the case of the sulfur purple bacteria, significant advances can only be expected from pure culture studies under controlled environmental conditions. Key to the genera of family Chlorobacteriaceae. I. Free-living bacteria not intimately associated with other microbes, a. Bacteria not united into well defined colonies. Genus I. Chlorobium, p. 869. aa. Bacteria united into characteristic aggregates. b. Bacteria without intracellular sulfur globules. Genus II. Pelodictyon, p. 870. bb. Bacteria with intracellular sulfur globules. Genus III. Clathrochloris, p. 872. II. Green bacteria found as symbiotic aggregates with other organisms, a. Aggregates composed of green bacteria and protozoa. Genus IV. Chlorobacteriuin, p. 872. aa. Aggregates composed of two different types of bacteria. b. Aggregates small, barrel-shaped, actively motile, and consisting of a central, polarly flagellated, rod-shaped bacterium with a covering of green sulfur bacteria. Genus V. Chlorochromatium, p. 873. bb. Aggregates large, cylindrical, non-motile, and composed of a central filamentous bacterium with a more or less extensive covering of green sulfur bacteria. Genus VI. Cylindrugloea, p. 873. Genus I. Chlorobium Nadson. (Xadson, Bull. Jard. Imper. Botan., St. Petersb., 12, 1912, 64 (Russian), 83 (Ger- man) ; Chloronostoc Pascher, Die Siisswasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 456; Tetrachloris Pascher, ibid., 455; Sorochloris? Pascher, * Completely revised by Prof. C. B. van Niel, Hopkins Marine Station, Pacific Grove, California, January, 1944. 870 MANUAL OF DETERMINATIVE BACTERIOLOGY ibid., 455; Chloropseudomonas? Czurda and Maresch, Arch. f. Mikrobiol., 8, 1937, 123; in part, Pelogloea Lauterborn, Verhandl. naturhistor.-medizin. Vereins, Heidel- berg, N.F. 13, 1915, 430.) From Greek Mows, green and bios, life. Green sulfur bacteria, occurring singly or in chains, individual cells of various sizes and shapes, from spherical to relatively long rod-shaped, the latter sometimes coiled into tight spirals; often united in chains, and generally embedded in a slime capsule. Non-motile. Contain a chlorophyllous pigment diiferent from the com- mon green plant chlorophylls and from bacterio-chlorophyll. Capable of photosyn- thesis in the presence of hydrogen sulfide, during which they produce elementary sulfur which is excreted outside the cells. Do not form spores. The type species is Chlorobium limicola Nadson. 1. Chlorobium limicola Nadson. (Nadson, Bull. Jard. Imper. Botan., St. P^tersb., 12, 1912, 64 (Russian), S3 (German) ; Chloronostoc abbreviation Pas- cher, Siisswasserfiora Deutschlands, Osterreichs und der Schweiz, Jena, 13, 1925, 456; Tctrachloris inconstans Pascher, ibid., 456; SorocJdoris aggre- gata? Pascher, ibid., 455; in part Pelo- gloea chlorina Lauterborn, Verhandl. naturhistor.-medizin. Vereins, Heidel- berg, N.F. 13, 1915, 430.) From Latin, mud-dweller. Cells: Various shapes and sizes, mark- edly dependent upon environmental conditions. In young and healthy state predominantly spherical to ovoid, about 0.5 to 1 micron in diameter, frequently united in chains resembling streptococci. Often cells become elongated and appear as rods, generally about 0.7 micron by 1 to 2.5 microns; also these may remain united in chains. Regularly produce mucus, causing the formation of cell- conglomerates of different size and shape, but not, as a rule, of characteristic appearance. Color yellowish-green. Non-motile. Abnormal cell forms (involution forms) rather common. These may be larger spherical cells, up to 5 to 6 microns in diameter, the larger ones generally vacuolated, or long rods, occasionally club-shaped but more often coiled. In rare cases the latter may be loosely wound. More frequently they are tightly-coiled screws, with cells of about 0.5 micron in diameter by as much as 15 microns in length. The spherical in- volution forms are normally encountered in acid, the coiled ones in alkaline en- vironments. Strictly anaerolnc and apparently dependent upon hydrogen sulfide and light. Development in organic media has not been obtained. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light; more rarely in sulfur springs. Illustrations: Nadson, loc. cit., PI. Ill, fig. 3-12; van Niel, Arch. f. Mikrobiol., 3, 1931, 66, fig. 8. Genus II. Pelodictyon Lauterborn. (Lauterborn, AUgem. botan. Ztschr., 19, 1913, 98; Verhandl. naturhistor.-medizin. Vereins, Heidelberg, N.F. 13, 1915, 431; Schmidlea loc. cit., Lauterborn, Allgem. botan. Zeitschr., 19, 1913, 97; in part, Pelogloea Lauterborn, Verhandl. naturhist. medizin Vereins, Heidelberg, N.F. 13, 1915, 430; Pediochloris Geitler, Die Susswasser- flora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 457.) From Greek pelos, mud and dictyon, net. Green sulfur bacteria, individual cells ovoid to distinctly rod-shaped, producing rather extensive mucoid capsules, and generally united into large colonies of char- acteristic shapes. Non-motile. Contain chlorophyllous pigments different from Family chlorobacteriaceae 871 the common green plant chlorophylls and from bacteriochlorophyll. Capable ol" photosynthesis in the presence of hydrogen sulfide, but do not store sulfur globules inside the cells. The type species is Pelodictyon clathr at i forme (Szafer) Lauterborn. Key to the species of genus Pelodictyon. I. Cells united in colonies in a net-like fashion. 1. Pelodictyon. clathralifonnc. II. Cells arranged in tightly packed colonies without net-like structure. a. Colonies composed of irregularly arranged cell-masses, e.xtending in three dimensions. 2. Pelodictyon aggregatum. aa. Colonies consisting of parallel strands and extending in two dimensions. 3. Pelodictyon parallelion. 1. Pelodictyon clathratiforme (Szafer) Lauterborn. {Aphanothece clathrati- forme Szafer, Bull. Acad. Sci., Cracovic, S6r. B,3, 1910, 162; Lauterborn, Allgem. botan. Ztschr., 19, 1913, 98; Lauterborn, Verhandl. naturhist.-medizin. Vereins, Heidelberg, N.F. 13, 1915, 430; Pelodic- tyon clathratiforme Geitler, Die Siiss- wasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 458; Pelodictyon lauterbornn Geitler, ibid., 458.) From Greek clathros, trellis and formis, shape. Cells: Generally rod-shaped, ranging from slightly elongated ovoids to dis- tinct rods, often vacuolated, about 0.5 to 1.5 micron by 2 to 4 microns, produc- ing rather wide slime capsules, and characteristically united into three- dimensional colonies which present a net-like appearance, with mazes of about 10 to 50 microns. Color yellowish-green. Xon-motile. Abnormal cell forms (involution forms) not uncommon, consisting of elongated and curved, forked, or club- shaped and swollen rods, occasionally suggesting rudimentary branching at the extremities. Such cells may be found as elements in chains for the greater part composed of normal individuals. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light; sulfur springs. Illustrations: Szafer, loc. cit., PI. VI, fig. 5; Perfiliev, Jour. Microbiol. (Rus- sian), 1. 1914. PI. II, fig. 1, 5-12; Lauter- born, loc. cit.. 1915, PI. Ill, fig. 33. 2. Pelodictyon aggregatum Perfiliev. {Aphanothece lutecla Schmidle, Beihefte Botan. Cent., 10, 1901, 179; Schmidlea luteola Lauterborn, Allgem. l)otan. Ztschr., 19, 1913, 97; Lauterborn, Ver- handl. naturhistor.-medizin. Vereins, Heidelberg, X.F., IS, 1915, 429; Pelo- gloea hacillifera Lauterborn, ibid., 430; Perfiliev, Jour. Microbiol. (Russian), i, 1914, 197.) From Latin aggregatus, leading together, grouping. Cells: Usually rod-shaped, about 1 to 1.5 microns by 2 to 4 microns, often vacuolated, producing mucus capsules, and united into irregularly shaped, three-dimensional colonies in which the cells are more or less tightly packed, without orderly arrangement. Colonies may attain a size of up to 1 mm; fre- quently they are not fully compact, but contain less dense areas, or appear per- forated, thus forming transition stages to Pelodictyon clathratiforme. Color yellowish-green. Non-motile. Abnormal cell forms (involution forms) usually in the shape of elongated and curved, forked or club-shaped and swollen rods, occasionally suggesting branching at extremities. Habitat : Alud and stagnant water, con- taining rather high concentrations o£ 872 MANUAL OF DETERMINATIVE BACTERIOLOGY hydrogen sulfide, and exposed to light; sulfur springs. Illustrations: Perfiliev, loc. cit., PI. II, fig. 2; Lauterborn, loc. cit., PI. Ill, fig. 29-31. 3. Pelodictyon parallelum Perfiliev. (Aphanothece parallela Szafer, Bull. Acad. Sci., Cracovie, S^r. B, 3, 1910, 163; Perfiliev, Jour. Microbiol. (Russian), 1, 1914, 198; Pediochloris parallela Geitler, Die Siisswasserflora Deutschlands, Os- terreichs und der Schweiz, Jena, 12, 1925, 457.) From Latin parallelus, beside one another. Cells: Rather small, spherical to ovoid, or even rod-shaped; about 0.5 to 1 micron by 1 to 3 microns, occurring in chains, and forming flat, plate-like, two-dimen- sional aggregates in which the chains are arranged as parallel strands. Color yellowish-green. Xon-motile. Abnormal cell forms not specifically mentioned, but likely to occur, and to resemble those of other species. This species may well be a special growth-form of Chlorobium limicola. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light; sulfur springs. Illustrations: Szafer, loc. cit., PI. VI, fig. 7; Perfiliev, loc. cit., PI. II, fig. 2. Genus III. Clathrochloris Geitler. (Die Susswasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 457.) From Greek clathros, trellis and chloros, green. Green sulfur bacteria of small size, generally spherical, and arranged in chains which are united into loose, trellis-shaped aggregates, somewhat similar to those of Pelodictyon clathratifornie and Pelodictyon aggregatum. Cells usually contain sulfur globules. Color yellowish-green. Non-motile. The type species is Clathrochloris snlphurica (Szafer) Geitler. 1. Clathrochloris sulphurica (Szafer) Geitler. {Aphanothece sulphurica Sza- fer, Bull. Acad. Sci., Cracovie, S6r.B, 3, 1910, 162; Geitler, Die Siisswasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 457.) From Latin, containing sulfur. Cells: Spherical, about 0.5 to 0.7 mi- cron in diameter, usually containing sulfur globules. Color yellowish-green. Xon-motile. The reported occurrence of sulfur globules in the cells of this very small species is surprising; it is the only one among the green sulfur bacteria in which these inclusions have been encountered. The published descriptions are even more fragmentary than those of other mem- bers of the group. Source: Reported only from sulfur .springs in Lubien Wielki, near Lwow, Poland. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light; sulfur springs. Illustration: Szafer, loc. cit., PI. VI, fig. 6. Genus IV. Chlorobacterixun Lauterborn. (Lauterborn, Verhandl. naturhist.-medizin. Vereins, Heidelberg, N.F., 13, 1915, 429; not Chlorobacterium Guillebeau, Landw. Landw. Jahrb. d. Schweiz, 4, 1890, 32; Chroostipes Pascher, Die Siisswasserflora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 116.) From Greek chloros, green and Latin bacterium, a small rod. Green sulfur bacteria(?) which grow symbiotically as an outside covering on cells of protozoa, such as amoeba and flagellates. Cells rod -shaped, often slightly curved, greenish. Non-motile. The type species is Chlorobacterium symbiolicum Lauterborn. FAMILY CHLOROBACTERIACEAE 873 1. Chlorobacterium symbioticum Lau- terborn. (Lauterborn, Verhandl. na- turhist.-medizin. Vereins, Heidelberg, N.F., 13, 1915, 429; Chroostipes linearis Pascher, Die Siisswasserflora Deutsch- lands, Osterreichs und der Schweiz, Jena, 13, 1925, 116.) From Greek, living symbiotically. Cells: Rod-shaped, about 0.5 by 2 to 5 microns, often slightly curved. Xon- motile. Occur as a peripheral covering of cer- tain protozoa with which they may form a symbiotic unit. It is iKjt certain that this is a green sulfur bacteriimi; the description of localities where it was found fail to mention the presence of hydrogen sul- fide in the environment which should be a prcM-equisite for a member of this group. Source: Reported from a number of ])ools in Germany. Habitat: Stagnant water. Illustrations: Lauterborn, loc. oil., PI. Ill, fig. 34-36; Pascher, loc. cit., fig. 149. Genus V. Chlorochromatiiun Laiderburn. (Lauterborn, Allgem. botan. Ztschr., 19, 1906, 196; Chloronium Buder, Ber. d. deut. bot. Ges., 31, 1914, General versammlungsheft, 80.) From Greek chloros, green and chroma, color. Green sulfur bacteria, ovoid to rod-shaped with rounded ends, occurring as barrel- shaped aggregates, consisting of a rather large colorless bacterium with a polar flagel- lum as the center, surrounded by the green bacteria, arranged in 4 to 6 rows, ordi- narily from 2 to 4 cells high. The entire conglomerate behaves like a unit, is motile, and multiplies by the more or less simultaneous fission of its components. The green constituents contain a chlorophyllous pigment which is not identical with the common green plant chlorophylls or with bacteriochlorophyll. Capable of photosynthesis in the presence of hydrogen sulfide, but do not store sulfur globules in the cells. The type species is Chlorochromaiium aggrcgatum Lauterborn. 1. Chlorochromatium aggregatum Lau- terborn. (Lauterborn, Allgem. botan. Ztschr., 19, 1906, 196; Chloronium mira- bile Buder, Ber. deut. botan. Ges., 31, 1914, Generalversammlungshef t , SO.) From Latin aggregatus, grouped. Cells of the green component 0.5 to 1.0 by 1.0 to 2.5 microns, mostly from 8 to 16 individuals surrounding the central bacterium. Size of the total barrel- shaped unit variable, generally 2.5 to 5 by 7 to 12 microns. Occasionally a group of the complex colonies may remain at- tached in a chain. Anaerobic. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light. There is at present no good reason for distinguishing 2 varieties (forma typica and forma minor) or even species, on the basis of size differences of the colonj-, as Geitler proposed (Die Siisswasser- flora Deutschlands, Osterreichs und der Schweiz, Jena, 12, 1925, 460). The re- portetl and personally observed sizes of such units show that the extreme limits are linked by a complete series of transi- tions. Illustrations: Buder, loc. cit., PI. XXIV, fig. 1-5; Perfiliev, Jour. Micro- biol. (Russian), 1. 1914, 213, fig. 1-5. (Jour. Microbiol. Greek gloios, a glutinous substance. Genus VI. Cylindrogloea Perfiliev. (Russian), /, 1914, 223.) From Latin cylindrus, cylinder and 874 MANUAL OF DETERMINATIVE BACTERIOLOGY Green sulfur bacteria, consisting of small ovoid to rod-shaped cells, growing in association with a filamentous, colorless, central bacterium, thus forming colonies of a cylindrical shape. Non-motile. The green component contains a chlorophyl- lous pigment different from the common chlorophylls of green plants and from bac- teriochlorophyll. Capable of photosynthesis in the presence of hydrogen sulfide, without depositing sulfur globules in the cells. The type species is Cylindrogloca bactcrifera Perfiliev. 1. Cylindrogloea bacterifera Perliliev. (Jour . Microbiol. (Russian) , 1 , 1914, 223.) From Latin bacter, rod and fero, to bear. Individual green components ovoid to rod-shaped, about 0.5 to 1 by 2 to 4 mi- crons, very similar to those of the com- plex Chlorobacleri am symhioticum and Chlorochromatium aggregatani with which they may well be identical. The central filamentous bacterium is embedded in a slime capsule of considerable dimensions. This, in turn, is surrounded by a layer of green bacteria, usually one cell thick. The green organisms may form a verj^ dense outer covering, or they may be more sparsely distributed over the mu- cus capsule. The entire unit is again surrounded by a sizeable slime zone. Aggregates measure about 7 to 8 microns in width, and up to 50 microns in length; they are non-motile. Both components appear to be non-spore-forming. Habitat: Mud and stagnant water con- taining rather high concentrations of hydrogen sulfide and exposed to light. Illustration: Perfiliev, loc. cit., 213, fig. 6-11. Perfiliev rightly emphasizes, as Buder had done for Chloronium mirabile, the provisional nature of thus using a gen- eric designation for an apparently stable complex composed of two different or- ganisms. It remains possible that the last three genera of symbiotic entities represent fortuitous combinations whose occurrence is conditioned by environ- mental factors. If so, the generic ter- minology would be devoid of any taxo- nomic significance, and the green bacteria should be relegated to more appropriate genera. Indications sugges- tive of this state of affairs can be found in the literature; for example in Uter- mohl's ob.servation (Archivf.H,ydrobiol., Suppl. 5, 1925, 279) that the complex Chlorochromalium aggregalum may, espe- cially in the presence of oxygen, dis- integrate, whereupon the green con- stituents appear as small Pelodictyon aggregalum (Schmidlea luteola) colonies. FAMILY MYCOBACTERIACEAE 875 ORDER II. ACTINOMYCETALES BUCHANAN. (Jour. Bact., 2, 1917, 162.) Organisms forming elongated cells which have a definite tendency to branch. These hyphae do not exceed 1.5 microns and are mostly about 1 micron or less in diameter. In the Mycobacteriaceae the mj'celium is rudimentary or absent ; no spores are formed ; the cells are acid-fast. The Aciinomycetaceae and Strepiomycelaceae usually produce a characteristic branching mycelium and multiply by means of special spores, oidio- .spores or conidia. Special spores are formed by fragmentation of the plasma within straight or spiral-shaped spore-bearing hj'phae; the oidiospores are formed by seg- mentation, or hy transverse division of hyphae, similar to the formation of oidia among the true fungi ; the conidia are produced singly, at the end of simple or branch- ing conidiophores. They grow readily on artificial media and form well-developed colonies. The surface of the colony, especially in the Aciinomycetaceae and Strepto- mijcetaceae , may become covered with an aerial mycelium. Some form colorless or white colonies, whereas others form a variety of pigments. Some species are par- tially acid-fast. In relation to temperature, most are mesophilic, while some are thermophilic. Certain forms are capable of growing at low oxj'gen tension. The Order as a whole is composed of saprophj'tic species, but also includes species that are parasitic and sometimes pathogenic on both animals and plants. Key to the families of order Actinomycetales. I. Mycelium rudimentary or absent, no spores formed. Acid-fast. Family I. Mycobacteriaceae, p S75. II. True mycelium produced. A. Vegetative mycelium divides by segmentation into bacillary or coccoid elements. Some species partially acid-fast. Family II. Actinomycetaceae, p. 892. B. Vegetative mj'celium normally remains undivided. Family III. Streptomycetaceae, p. 929. Among the recent sj'stems of classification of this order it is sufficient to mention the following: Baldacci (INR-copath., 2, 1939, 84) divided the order Actinomycetales into two families: (a) Mycobacteriaceae Chester with two subfamilies, Leptotrichi- oideae Baldacci and Proactiiiomycoideae Baldacci, each with five genera, and (b) Actinomycetaceae Buchanan, with two genera, Micromonospora and Actinomyces. Krassiluikov (Ray fungi and related organisms, Izd. Akad. Nauk, Moskow, 1938) divided the order into (a) Actinomycetaceae, with four genera, Actinomyces, Proac- tinomyces, Mycobacterium and Mycococcus, and (b) Mtcromonosporaceae, with one genus, Micromonospora. Waksman (Jour. Bact., 39, 1940, 549) divided the order into four families: Mycobacteriaceae, Proactinomycetaceae, Actinomycetaceae and Mtcro- monosporaceae. FMIILY I. >IYCOBACTERIACEAE CHESTER.* (Chester, Man. Determ. Bact., 1901, 349; Proactinomycetaceae Lehmann and Haag, in Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 674.) * Completeh' revised by Prof. G. B. Reed, Queens University, Kingston, Ontario, Canada, December, 1938; minor revisions, December, 1944; with a complete revision of Mycobacterium, leprae and ]\l. Icpracmurium by Dr. John H. Hanks, Leonard Wood Memorial, American Leprosy Foundation, New York, N. Y. 876 MANUAL OF DETERMINATIVE BACTERIOLOGY Slender filaments, straight or slightly curved rods, frequently irregular in form with only slight and occasional branching. Often stain unevenly, i.e., show variations in staining reaction within the cell (beading). Xo conidia. Non-motile. Aerobic. Gram-positive. Acid-fast. t Pathogenic species grow slowly (several weeks); those from soil, water and vegetation more rapidly (several days). There is a single genus Mycobacterium Lehmann and Neumann. Genus I. Mycobacterium Lehmann and Neumann. {Coccothrix Lutz, Zur Morphologic des Mikroorganismus der Lepra. Dermatolo- gische Studien, Heft 1, 1886, 22; Schleroihrix Metschnikoff, Arch. f. path. Anat. u. Physiol., 113, 1888, 70; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 108; Mrjcomonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 329; Eumrjces Battaglia, Soc. Internaz. Microbiol. Boll. Sez. Ital., 10, 1938, 166.) From Greek myces, fungus and bacterium, a little rod. Characters as for the family. The type species is Mycobacterium tuberculosis (Schroeter) Lehmann and Neumann. Key to the species of t/enus Mycobacterium. I. Parasites in warm-blooded animals; grow slowly on all media. A. Grow slowly on glycerol agar in atmospheric air; experimentally infect guinea pigs and fowls. 1. Experimentally produces generalized tuberculosis in guinea pigs but not in rabbits and fowls. Growth enhanced by the addition of glycerol to most media. Generally pale yellow to orange pigmenta- tion on serum media. la. Mycobacterium tuberculosis var. hominis. 2. Experimentally produces generalized tuberculosis in guinea pigs and rabbits but not in fowls. Growth not enhanced by addition of glycerol to media. Never pigmented. lb. Mycobacterium, tuberculosis var. bovis. 3. Experimentally produces generalized tuberculosis in fowls and rab- bits but not in guinea pigs. 2. Miicobadcrium avium. B. Grows in primary culture on glycerol agar only when extracts of, or heat- killed acid-fast bacilli are added. Experimentally fails to infect guinea pigs or fowls. 3. Mycobacterium, paraiuberculosis. C. Have not been grown on culture media thus far devised. Exi)erimentally fail to infect guinea pigs or fowls. 1. Has not experimentally been transmitted to any animal species. 4. Mycobacterium leprae. 2. Occurs in wild rats, and can be experimentally transmitted to rats and some strains of mice. 5. Mycobacterium lepraemurium. t Most acid-fast bacteria treated with carbol-auramin and decolorized with NaCl- HCl-alcohol show fluorescence under the microscope when they are radiated by long wavelength ultraviolet light (Haitinger, Fluorescenz-mikroskopie, Leipzig, 1938, 108 pp.; Ellinger, Biol. Revs., 15, 1940, 323-350; Richards, Jour. Bact., U, 1942, 721). For a discussion of the influence of environment on acid-fastness, see Salle and Moser, Internat. Jour. Leprosy, 5, 1937, 163. FAMILY MYCOBACTERIACEAE 877 II. Saprophytes or parasites on cold-blooded animals ; grow rapidly on most media. A. Fail to survive 60°C for 1 hour. 1. Fail to grow at 47°C. a. Unable to utilize sorbitol. 6. Mycobacterium piscium. 7. Mycobacterium marinum, 8. Mycobacterium ranae. 9. Mycobacterium thamnopheos. aa. Utilize sorbitol. 2. Grows at 47°C. 10. Mycobacteriu?n friedmannii. 11. Mycobacterium spp. 12. Mycobacterium lacticola. Survives 60°C for 1 hour; grows at 47°C. 13. Mycobacterium phlei. 1. Mycobacterium tuberculosis (Schroeter) Lehmann and Neumann. (Tuberkelbacillen, Koch, Mitteil. a.d. kaiserlich. Gesundheitsamte, 2, 1884, 6; Bacillus tuberculosis Schroeter, in Cohn, Kryptogamen Flora v. Schlesien, 3, 1886, 164; Bacillus tuberculosis Fliigge, Die Mikroorganismen, 2 Aufi., 1886, 208; Coccothrix tuberculosis Lutz, Dermatol. Studien, 1, 1886, 22; Sclerothrix kochii Metchnikoff, Arch. f. path. Anat. u. Physiol., 113, 1888, 70; Bacterium tuber- culosis Migula, in Engler and Prantl, Die nattirlichen Pflanzenfamilien, I Abt., la, 1895, 23; Lehmann and Neu- mann, Bakt. Diag., 1 Aufi., 2, 1896, 363; Bacillus kochii, quoted from Lehmann and Neumann, idem; Discomyces tubcrcu- losus (sic), quoted from Neveu-Lemaire, Precis de Parasitol. Humaine, 5th ed., 1921, 25; Sclerothrix tuberculosis Vuille- min,Encyclopedie Mycologique, Paris, ^, 1931, lM;Eu7nyces tuberculosis Battaglia, Soc. Internaz. Microbiol. Boll. Sez. Ital., 10, 1938, 166.) From M. L. tuber- culosis, tuberculosis. Two varieties of this species are com- monly recognized, the human and the bovine. la. Mycobacterium tuberculosis var. hominis Lehmann and Neumann. (Hu- man tubercle bacilli, Th. Smith, Trans. Assoc. Am. Phys., 11, 1896, 75; Myco- bacterium tuberculosis typus humanus Lehmann and Neumann, Bakt. Diag., 4 Aufi., 2, 1907, 550.) From Latin hominis, of man. Common name : Human tubercle ba- cillus. Description from Koch (loc. cit.) and Topley and Wilson (Princip. of Bact. and Immun., London, 2nd ed., 1936, 315). Rods, ranging in size from 0.3 to 0.6 by 0.5 to 4.0 microns, straight or slightly curved, occurring singly and in occa- sional threads. Sometimes swollen, cla- vate or even branched. Stain uniformly or irregularly, showing banded or beaded forms. Acid-fast and acid-alcohol -fast. Gram-positive. Growth in all media is slow, requiring several weeks for de- velopment . This bacterium contains mycolic acid (Stodola, Lesuk, and Anderson, Jour. Biol. Chem., 136, 1938, 505-513). The acid-fast mycolic acid combines more firmly with carbol-auramin than with carbol-fuchsin and this apparently ac- counts for the increased sensitivity of fluorescence microscopy for this bac- terium (Richards, Science, 93, 1941, 190; Richards, Kline, and Leach, Amer. Rev. Tuberc, U, 1941, 255-266). Nutrient agar: No growth. Glycerol agar colonies : Raised, thick, cream-colored, with a nodular or wrinkled surface and irregular thin margin. Glycerol agar slant: After 4 weeks, raised, thick, confluent, cream-colored gro^'th. Nutrient broth : No growth. 878 MANUAL OF DETERMIXATIVE BACTERIOLOGY Glycerol broth: After 8 weeks, thick, white or cream-colored, wrinkled pellicle extending up the sides of the flask, no turbidity; granular or scaly deposit. Dorset's egg slants: After 4 weeks, rather sparse, discrete or confluent, slightly raised, grayish-yellow growth with finely granular surface. Glycerol egg slants : After 4 weeks, luxuriant, raised, confluent, gray to yellow growth, with granular surface, generally with nodular heaped-up areas. Coagulated beef serum: After 4 weeks, thin, effuse, confluent, gray to yellow growth, with a very fine granular surface. Gl3^cerol beef serum: After 4 weeks, luxuriant, thick, raised, confluent, yellow to orange -yellow growth, with coarsely granular surface, generally with irregu- larly heaped-up areas. Litmus milk : Growth, but no change in the milk. Glycerol potato: After 4 weeks, luxu- riant, raised, confluent, cream-colored growth with a nodular or warty surface. Carbohydrates : Glucose, fructose, arab- inose and galactose are utilized ; sucrose and lactose not utilized (Merrill, Jour. Bact., SO, 1930, 235, based on the exam- ination of one strain). Optimum temperature 37 °C. Optimum pH 7.4 to 8.0 (Ishimori, Ztschr. f. liyg., 102, 1924, 329); pH 6.0 to 6.5 (Dernby and Naslund, Biochem. Zeit., 133, 1922, 392). Pathogenicity : Produces tuberculosis in man, monkey, dog and parrot. Ex- perimentally, it is highly pathogenic for guinea pigs but not for rabbits, cats, goats, oxen or domestic fowls. Intermediate manmialian types : Griffith (Lancet, 1, 1916-17, 721; Jour. Path, and Bact., 21, 1924, 54) has found aberrant types particularly in skin lesions of both man and ox, which are in certain characteristics intermediate between the human and the bovine varieties. He finds no evidence, however, that the one variety may change into the other. Variation: Variation in colony struc- ture of the two mammalian varieties. comparable with that in other species, has been described by several authors, as Petroff et al. (Jour. E.xp. Med., 60, 1934, 515), Birkhaug (Ann. Inst. Past., 57, 1933, 428), Kahn et al. (Jour. Bact., 25, 1933, 157), Uhlenhuth and Sieffert (Zeit. Immun., 59, 1930, 187), Reed and Rice (Canad. Jour. Res., 5, 1931, 111), Smith- burn (Jour. Exp. Med., 63, 1936, 95) and Shaffer (Jour. Path, and Bact., 40, 1935, 107). Several of these authors have found associated variation in cell struc- ture and in virulence though Boquet (Compt. rend. Soc. Biol. Paris, 103, 1930, 290), Birkhaug (Ann. Inst. Past., 49, 1932, 630), and others, have failed to find differences in virulence. Reed and Rice (Jour. Immunol., 23, 1932, 385) found the S form to contain an antigenic substance lacking in the R form. Antigenic structure: By agglutination, absorption of agglutinins and comple- ment fixation a distinction may be made between the mammalian varieties and Mycobacterium avium, but it has been im- possible to distinguish, by these means, between the two mammalian varieties (Tullock et al.. Tubercle, 6, Oct.-Dec, 1924, 18, 57 and 105; Wilson, Jour. Path, and Bact., 28, 1925, 69; Griffith, Tubercle, 6, May, 1925, 417; Rice and Reed, Jour. Immunol., 23, 1932, 385; Kauffman, Ztschr. f. Hyg., 114, 1932, 121). Tu- berculins prepared from the human and the bovine varieties are ordinarily in- distinguishable in their action but Lewis and Seibert (Jour. Immunol., 20, 1931, 201) detected a difference bj' cross an- aphylactic reactions. Distinctive characters : Tubercle ba- cilli pathogenic for guinea pigs and rab- bits, not for fowls. Mycobacteriiun tuberculosis var. ho7ninis produces gen- eralized tuberculosis in guinea pigs but not in rabbits. Mycobacterium tubercu- losis var. bovis produces generalized disease in both guinea pigs and rabbits. Growth of the human variety is enhanced by the addition of glj^cerol to most media. The growth of the bovine variety is not enhanced by the addition FAMILY MYCOBACTERIACEAE 879 of glycerol. The human variety gen- erally develops yellow to red pigment on serum media, while the bovine variety never produces pigment. Antigenically the two varieties are not distinguishable. Source : From tuberculous lesions in man. Habitat : The cause of tuberculosis in man. Transmissible to rabbits and guinea pigs. lb. Mycohacterium tuberculosis var. bovis Lehmann and Neumann. (Bovine tubercle bacilli, Th. Smith, Trans. Assoc. Am. Phys., 11, 1896, 75; 13, 1898, 417; Jour. Exp. Med., 3, 1898, 451; M)jcobacterium tuberculosis typus bo- vimis Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 550.) From Latin bovis, of the ox. Common name : Bovine tubercle ba- cillus. Description from Th. Smith {loc. cit.) and Topley and Wilson (Princip. of Bact. and Immun., 2nd ed., 1936, 315). Rods which are shorter and plumper than the human type. Range in size from 1.0 to 1.5 microns. Very short forms are frequently intermixed with somewhat larger forms. Stain regularly or irregularly. Acid-fast and acid-alco- hol-fast. Gram-positive. Less easily cultivated than the human variety. Nutrient agar : No growth. Glycerol agar colonies : Small, irregular, with granular surface, no pigment. Glycerol agar slant: After 4 weeks, thin, granular or effuse, confluent growth. Nutrient broth: No growth. Glycerol broth: After 8 weeks, thin grayish-white film, slightly nodular, no turbidity. Slight granular deposit. Dorset's egg slants: After 4 weeks, similar to var. hominis but generally poorer growth and no pigmentation. Glycerol egg slants: After 4 weeks, similar to Dorset's egg slants. Coagulated beef serum: After 4 weeks, thin, effuse, confluent, white to gray growth with very fine granular surface. Generally less luxuriant than in the human variety. Glycerol beef serum: After 4 weeks, similar to plain beef serum. Glycerol potato : After 4 weeks, thin, effuse, grayish growth. Litmus milk : Growth, but no change in the milk. Optimum temperature 37°C. Optimum pH 5.8 to 6.9 (Ishimori, Ztschr. f. Hyg., 102, 1924, 329) ; 6.0 to 6.5 (Dernby and Naslund, Biochem. Zeit., 132, 1922, 392). Pathogenicity : Produces tuberculosis in ox, man, monkey, goat, sheep, pig, cat, parrot, cockatoo and possibly some birds of prey. Experimentally, it is highly pathogenic for rabbit and guinea pig, slightly pathogenic for dog, horse, rat and mouse; not pathogenic for fowls. Variation: See Mycobacterium tubercu- losis var. hominis. Antigenic structure : See Mycobacte- rium tuberculosis var. hominis. Distinctive characters : See Mycobac- terium tuberculosis var. hominis. Source: From tubercles in cattle. Habitat : The cause of tuberculosis in cattle. Transmissible to man and do- mestic animals. More highly pathogenic for animals than the human type. 2. Mycobacterium avium Chester. (Tuberculose des oiseaux, Strauss and Gamaleia, Arch. Med. exp. et Anat. path., 1891; Bacillus der Hiihnertu- berculose, Maffucci, Ztschr. f. Hygiene, 11, 1892, 449; Bacillus tuberculosis gal- linarum Sternberg, Man. of Bact., 1893, 392; Mycobacterium tuberculosis avium Lehmann and Neumann, Bakt. Diag., 1 Aufl., /, 1896, 370; Bacillus tuberculosis avium Kruse, in Fliigge, Die Mikro- organismen, 3 Aufl., 2, 1896, 506; Myco- bacterium avium Chester, Manual Determ. Bact., 1901, 357; Mycobacterium tuberculosis typus gallinaccus Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 553.) From Latin avis, bird. Common name : Avian tubercle ba- cillus. 880 MANUAL OF DETERMINATIVE BACTERIOLOGY Description from Strauss and Gamaleia (loc. cit.) and Topley and Wilson (Princip. of Bact. and Immun., 2nd ed., 1936, 315). Rods resembling those of the bovine type of tubercle organism. Nutrient agar: After 4 weeks, slight growth, effuse, translucent with fine granular surface. Glycerol agar colonies : After 3 to 4 weeks, raised, regular, hemispherical, creamy or white colonies. Nutrient broth: After 4 weeks, very slight viscous to granular bottom growth, no pellicle, no turbidity. Glycerol broth: After 4 weeks, diffuse, turbid growth with a viscous to granular deposit. Dorset's egg slants : After 4 weeks, con- fluent, slightly raised growth, with smooth regular surface. Glj'cerol egg slants: After 4 weeks, luxuriant, raised, confluent, creamy to yellow growth with perfectly smooth surface. Coagulated beef serum : After 4 weeks, thin, effuse, grayish-yellow growth with smooth surface. Glycerol beef serum: After 4 weeks, luxuriant, raised, confluent, yellow to orange-yellow or occasionally pale pink growth, with a smooth glistening surface. Glycerol potato: After 4 weeks, luxu- riant, raised, confluent, with smooth to nodular surface. Litmus milk : Growth, but no change in the milk. Carbohydrates : Fructose, arabinoseand sucrose are utilized, glucose is slightly utilized, galactose and lactose are not utilized (Merrill, Jour. Bact., 20, 1930, 235, based on the e.xamination of one strain). Optimum temperature 40°C ; range 30° to 44°C (Bynoe, Thesis, McGill Uni- versity, Montreal, 1931). Optimum pH 6.8 to 7.3 (Bynoe, loc. cit.). Pathogenicity : Produces tuberculosis in domestic fowls and other birds. In pigs it produces localized and sometimes disseminated disease. Experimentally in the rabbit, guinea pig, rat and mouse it may proliferate without producing macroscopic tubercles — tuberculosis of the Yersin type. Man, ox, goat, cat, horse, dog and monkey are not infected. Variation : Winn and Petroff (Jour. Exp. Med., 57, 1933, 239), Kahn and Schwartzkopf (Jour. Bact., 25, 1933, 157), Birkhaug (Ann. Inst. Pasteur, 5Jt, 1935, 19), Reed and Rice (Canad. Jour. Res., 5, 1931, 111) and others, have shown variation to follow the course described for many species. Winn and Petroff have separated four colonial types : smooth, flat smooth, rough, deep yellow smooth. These also differ in chemical and physical properties. The smooth form exhibited the greatest degree of virulence, the flat smooth a lower viru- lence, while the chromogenic smooth and the rough were relatively benign. Some authors have failed to demonstrate this difference in virulence. The above de- scription applies primarily to the smooth form. Antigenic structure : By agglutination, absorption of agglutinins and comple- ment fixation Mycobacterium avium may be distinguished from other members of the genus (Tullock et al.. Tubercle, 6, 1924, 18, 57 and 105; Wilson, Jour. Path, and Bact., 28, 1925, 69; Mudd, Proc. Soc. Exp. Biol, and Med., 23, 1925, 569, and others). Furth (Jour. Immunol., 12, 1926, 273) and Shaffer (Jour. Path, and Bact., 40, 1935, 107) on this basis divided Mycohacterium avium into 1 or 2 sub- groups . Distinctive characters : Tubercle ba- cilli pathogenic for fowls, not for guinea pigs or rabbits . Culturally distinguished from the mammalian types by the ab- sence of pellicle formation in fluid media and the habit of growth on most solid media. Antigenically distinguished from other species. Source : From tubercles in fowls, widely distributed as the causal agent of tu- berculosis in birds and less frequently in pigs. FAMILY .MYCOB \CTEI{IACEAE 881 Habitat : The cause of tuberculosis in chickens. Transmissible to pigeon, other birds, mouse, rabbit and pig. 3. Mycobacterium paratuberculosis Bergey et al. (Darmtuberculose bacillen, Johneand Frothingham, Deutsch. Ztschr. Tiermed., 21, 1895, 438; Pseudotuberku- lose bacillen. Bang, Berl. tierarztl. Wchnschr., 1906, 759; Bacillus of Johne's Disease, M'Fadyean, Jour. Comp. Path., 20, 1907, 48; Twort, Proc. Roy. Soc, B, 83, 1910, 156; Bergey et al.. Manual, 1st ed., 1923, 374.) From M. L. -para- tuberculosis, of the disease paratubercu- losis. Common name: Johne's bacillus. The organism from a similar disease in sheep is probably identical though more difficult to cultivate 'Dunkin and Bal- four-Jones, Jour. Comp. Path., J^8, 1935, 236). Description from M'Fadj^ean {loc. cit.) and Twort and Ingram (A Monograph on Johne's Disease, London, 1913). Plump rods, 1.0 to 2.0 microns in length, staining uniformly, but occasionally the longer forms show alternately stained and unstained segments. Non -motile. Acid-fast. The organism is difficult to cultivate and, in primary cultures, has only been grown in media containing dead tubercle bacilli or other dead acid-fast bacteria (Boquet, Ann. Inst. Pasteur, 37, 1928, 495). In a few instances cultures have been acclimatized to a synthetic medium free from added dead bacteria (Dunkin, Jour. Comp. Path, and Therap., J^6, 1933, 159; Watson, Canad. Pub. Health Jour., 26, 1935, 268). Colonies on ghcerol agar containing heat-killed Mycobacterium phlei: After 4 to 6 weeks, just distinguishable, dull- white, raised, circular colonies. Colonies on Dorset's glycerol egg con- taining heat-killed Mycobacterium phlei: After 4 to 6 weeks, minute, dull-white, raised, circular, with a thin, slightly irregular margin. Older colonies become more raised, radially striated or irregu- larly folded and dull j-ellowish-white. Dorset's glycerol egg containing sheep's brain and heat-killed Mycobacterium phlei: Growth slightly more luxuriant. Glycerol broth containing heat -killed Mycobacterium phlei: Thin surface pel- licle which later becomes thickened and folded. Dorset's sj'nthetic fluid containing heat -killed Mycobacterium phlei: As on glj^cerol broth with M ycobacterium phlei. Pathogenicity: Produces Johne's dis- ease, chronic diarrhea, in cattle and sheep. Experimentally it produces a similar disease in bovine animals, sheep and goats. Guinea pigs, rabbits, rats and mice are not infected. Very large doses in laboratory animals produce slight nodular lesions comparable with those produced by Mycobacterium phlei. Antigenic structure : Johnin, prepared as tuberculin, gives positive reactions in cattle with Johne's disease. According to M'Fadyean et al. (Jour. Comp. Path, and Therap., 29, 1916, 62) tuberculous animals may also give a reaction. Plumb (Den Kong. Vet. Landboh0jskole Arssk., 1925, 63) has shown that a reac- tion maj' be produced in animals sensi- tized to avian tuberculin and that avian tuberculin causes a reaction in some ani- mals infected with Johne's bacillus. Distinctive characters : A small acid- fast bacillus producing characteristic lesions in cattle and growing only in the presence of dead acid-fast bacilli. Source : From the intestinal mucous membrane of cattle suffering from chronic diarrhea. Apparently an obligate para- site. Habitat : The cause of Johne's disease, a chronic diarrhea in cattle. The bac- teria are found in the intestinal mucosa. Xot pathogenic for guinea pigs or rabbits. 4. Mycobacterium leprae (Armauer- Hansen) Lehmann and Xeumann. (Ba- cillus leprae Armauer-Hansen, Norsk. Mag. laegevidensk., 9, 1874, 1; Arch, f. path. Anat. u. Physiol., 79, 1879, 882 MANUAL OF DETERMINATIVE BACTERIOLOGY 32; Nord. Med. Ark., 12, 1880, 1; Quart. Jour. Micro. Sci., 20, 1880, 92; Coccothrix leprae Lutz, Dermatol. Stud. 1, 1886, quoted from DeToni and Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 944; Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 372; Discomyces leprae Neveu-Lemaire, Precis Parasit. Hum., 5th ed., 1921, 27; Sclerothrix leprae Vuillemin, Encyclopedic Mj'co- logique, Paris, ^, 1921, 135; Mycobacterium leprae hominus Lowe, Internat. Jour. Leprosy, 5, 1937, 312.) From Greek lepra, leprosy. Common name: Leprosy bacillus. Armauer-Hansen Hoc. cit.) was the first to observe the bacilli in the tissues of lepers. The disease is now known as Hansen's disease. The bacilli occur in enormous numbers in lepromatous (nod- ular) cases of the disease and sparsely in the neural form. The present bac- teriological means of identification de- pend on: (a) acid-fast staining, and (b) failure of the organism to grow in bac- teriological media or in laboratory animals. Heated suspensions of the bacilli (obtained from nodules) produce a positive lepromin reaction in 75 to 97 per cent of normal persons and of neural cases of leprosy, but usually produce no reaction in lepromatous individuals (Mitsuda: See Hayashi, Int. Jour. Leprosy, /, 1933, 31-38). The failure of lepromatous persons to respond to in- jected leprosy bacilli constitutes a fundamental criterion for testing the validity of microorganisms such as other acid-fast or diphtheroid cultures which can at times be recovered from leprous tissues by inoculation of bacteriological media. Many organisms have been isolated from leprous tissues, some of which are acid-fast and have been styled Myco- bacterium leprae. The strains which have been adequately studied have proven to fall into the saprophytic groups (see No. 11, Mycobacterium, spp.) Hanks (Int. Jour. Leprosy, 9, 1941, 275- 298) found that acid-fast cultures of this type, as well as the diphtheroids which also have repeatedly been isolated from leprosj% were recoverable only from lesions located proximally with respect to open ulcers in the skin. Description of organisms seen in leprosy tissue from Armauer-Hansen {loc. cit.) and Topley and Wilson (Prin- cip. Bact. and Immun., 2nd ed., 1936, 316). Rods : 0.3 to 0.5 by 1 to 8 microns, with parallel sides and rounded ends, staining evenly or at times beaded. When nu- merous, as from lepromatous cases, they are generally arranged in clumps, rounded masses or in groups of bacilli side by side. Strongly acid-fast. Gram- positive. Pathogenicity : The communicability of leprosy from man to man is accepted (Rogers and Muir, Leprosy, 2nd ed., Baltimore, 1940, 260 pp.). Experimental transmission to humans or to animals has not been successful. Source: Human leprous lesions. In the lepromatous form of the disease bacilli are so abundant as to produce stuffed-cell granulomas; in the tubercu- loid and neutral lesions they are rare. Habitat: Obligate parasite in man. Confined largely to the skin (especially to convex and exposed surfaces) and to peripheral nerves. The microorganisms probably do not grow in the internal organs. 5. Mycobacteriun lepraemuriiun Mar- choux and Sorel. (Bacillus der Ratten- lepra, Stefanskj^, Cent. f. Bakt., I Abt., Orig., 33, 1903, 4S1 ; Mycobacterium leprae murium Marchoux and Sorel, Ann. Inst. Past., 26, 1912, 700; Bacillus leprae mu- rium Muir and Henderson, Indian Jour. Med. Res., 15, 1927, 15.) Mycobacterium pulviforme Marchoux (Ann. Derm., 1921, No. 21 and Ann. Inst. Past., 37, 1923, 348) from leprosy- like lesions in a man from Hayti is thought by the author to be identical with Mycobacterium lepraemurium. Common name: Rat leprosy bacillus. FAMILY MYCOBACTERIACEAE 883 Rods: 3.0 to 5.0 microns in length with slightly rounded ends. When stained, often show irregular appearance. Strongly acid-fast. Gram-positive. Like the human leprosy bacillus, this organism has not been cultivated in vitro; but can be passed experimentally through rats and some strains of mice. Distinctive features: The heat-killed bacilli produce lepromin reactions in lepratomous humans. The bacilli from lesions are not bound together in clumps, rounded masses and palisades as in hmnan lesions. For further details see review by Lowe (Internat. Jour. Leprosy, 5, 1937, 310 and 463). Source: An endemic disease of rats in various parts of the world, having been found in Odessa, Berlin, London, Xew South Wales, Hawaii, San Francisco and elsewhere. Habitat: The natural disease occurs chiefly in the skin and lymph nodes, causing induration, alopecia (loss of hair) and eventually ulceration. 6. Mycobacterium piscium Bergey et al. {Bacillus tuberculosis ■piscium Du- bard, Bull. acad. de med., 3 ser., 88, 1897, 580; Bataillon, Dubard and Terre, C!ompt. rend. Soc. Biol., 4, ser. 10, 1897, 446; Bergey et al., Manual, 1st ed., 1923, 375.) From Latin piscis, fish. Description from Bataillon et al. {loc. cit.) and Aronson (Jour. Inf. Dis., 39, 1926, 319). Slender rods, occurring singly and in threads, occasionally showing branching. Acid-fast. Non-motile. Gram-positive. Agar colonies : Small, circular, white, moist, with lobate margin and fine granular surface. Agar slant : Scant, white, moist, cream- like. Glycerol agar colonies: Thin, flat, smooth, glistening, yellow. Dorset's egg medium: Flat, smooth, moist, greenish. Broth: Thin pellicle, with flocculent sediment. Litmus milk: Thickened. No coagu- lation. Slightly alkaline. Potato: White, warty, butyrous colo- nies. Carbohydrates : Utilizes glucose and fructose but not sucrose, lactose, arabi- nose or galactose (IVIerrill, Jour. Bact., 20, 1930, 235, based on examination of one strain). Antigenic structure : By agglutination and complement fixation (Mudd, Proc. Exp. Biol, and Med., 23, 1925, 569; and Furth, Jour. Immunol., 12, 1926, 286) Mycobacterium piscium has been dis- tinguished from Mycobacterium fried- mannii, Mycobacterium ranae and prob- ably Mycobacterium marinum. From the limited number of cultures examined it is not evident whether this is due to species or strain specificity. Pathogenicity : Experimentally pro- duces tubercles in carp, frog and lizard, but not pathogenic for rabbit, guinea pig or birds (Dubard, Rev. de la Tuberc, 6, 1898, 13). Not pathogenic for salt water fish except eels (Betegh, Cent. f. Bakt., I Abt., Orig., 53, 1910, 374; 54, 1910, 211). Distinctive characters : Mycobacte- rium piscium, Mycobacterium marinum, Mycobacterium ranae, Mycobacterium thamnopheos and Mycobacterium fried- mannii constitute a closely related group — possibly one species . They differ from other members of the genus in their pathogenicity for cold-blooded animals, their failure to survive 60°C for an hour, their failure to grow at 47°C and their inability to utilize sorbitol. Mycobacterium marinum is distin- guished by its diffuse gro'v\i;h in broth, acid production in milk and deep yellow to orange pigmentation on most media. The other species grow in broth as a pellicle and render milk alkaline. Myco- bacterium piscium, Mycobacterium ranae, Mycobacterium thamnopheos and Myco- bacterium friedmannii may be distin- guished from each other by their habit of growth on solid media. But rela- tively few cultures have been studied 884 MANUAL OF DETERMINATIVE BACTERIOLOGY and the reports in certain important respects are conflicting, especially con- cerning pigmentation and utilization of carbohydrates. Aronson, Mudd and Furth found them to differ antigenically, but too few cultures were used to distin- guish between species and strain speci- ficity. Source : From tubercles in carp. Habitat : The cause of nodule and tu- mor-like formations in carp {Ciprinus carpio). Infectious for carp, frogs, liz- ards. Not infectious for guinea pigs and pigeons. 7. Mycobacterium marinum Aronson. (Jour. Inf. Dis., 39, 1926, 315.) From Latin marinus, marine. Description from Aronson {loc. cit.). In lesions, short, thick, uniformly staining organisms are seen frequently occurring in clumps, while long, thin, beaded or barred rods are scattered more discretely. In cultures the organisms have the same appearance. Non-motile. Acid-fast and acid -alcohol -fast. Gram- positive. Agar slant (slightly acid) : In five to seven days, moist, glistening, elevated colonies, becoming lemon-yellow. Gelatin: Not liquefied. Agar colonies : In 5 to 7 days, smooth, moist, slimy, lemon-yellow, later orange- colored. Glycerol agar colonies : In 14 to IS days, grayish-white, moist, elevated with irregular margins. Old growths lemon- yellow and still later orange -colored. Dorset's and Petroff's egg media: Similar to growth on glycerol agar but more luxuriant. Broth and glycerol broth : Growth is diffuse, no pellicle formed. Litmus milk : Acidified and coagulated. Indole not formed. Nitrites not produced from nitrates. Carbohydrates : Utilizes arabinose and fructose, fails to utilize sorbitol and galactose (Gordon, Jour. Bact., 84, 1937, 617). Aerobic, facultative. Optimum temperature 18° to 20°C. Fails to survive 60°C for 1 hour, fails to grow at 47°C (Gordon, Jour. Bact., 34, 1937,617). Pathogenicity : Experimentally infects salt water fish, goldfish, frogs, mice and pigeons, but not rabbits or guinea pigs. Antigenic structure : By agglutination and complement fixation distinguished from Mycobacterium ranae, Mycobacte- rium friedmannii, and probably Myco- bacterium piscium (Mudd, Proc. Soc. Exp. Biol, and Med., 23, 1925, 569 ; Furth, Jour. Immunol., 12, 1926, 286). See Mycobacterium piscium . Distinctive characters : See Mycobacte- rium piscium. Source : From areas of focal necrosis of the liver of sergeant majors (Abudefduf maurilii), croakers (Micropoyon undula- tus) and sea bass (Centropristes striatus). Habitat: Causes spontaneous tubercu- losis in salt water fish. 8. Mycobacterium ranae (Kuster) Bergey et al. (Kiister, Miinch. med. Wchnschr., 52, 1905, 57; Bergey et al., Manual, 1st ed., 1923, 374.) From Latin rana, frog. Description from Kuster {loc. cit.), Bynoe (Thesis, McGill University, Mon- treal, 1931) and Aronson (Jour. Inf. Dis., 44,1929,222). Slender rods, 0.3 to 0.5 by 2 to 8 mi- crons, smaller in old cultures. Uni- formly acid-fast in cultures 2 weeks old or older. In younger cultures the stain- ing is irregular, many organisms are not acid-fast. Non-motile. Gram-positive. Gelatin stab: No liquefaction. Agar colonies : Irregular, raised colo- nies, 1 to 3 mm in diameter with moist glistening surface, later becoming coarsely granular. Agar slant : Thick, spreading, glisten- ing. In old cultures dry and scaly. Putrid odor. Grayish-white. Glycerol agar colonies : Similar to gela- tin colonies but slightly creamy and be- coming dry and wrinkled in old cultures. FAMILY MYCOBACTERL\CEAE 885 Dorset's egg medium: Spreading, raised, glistening, later wrinkled. Loeffler's medium: Similar to Dorset's egg medium, white to buff-colored. Litmus milk: Becomes alkaline. Glycerol broth : Grayish flaky pellicle which breaks up early and settles. Broth: Slightly turbid, with slight sediment . Potato : Scanty, grayish growth, raised with a warty surface. Indole not formed. Nitrites are produced from nitrates. Carbohydrates: Glucose, fructose and arabinose are utilized; sucrose, lactose and galactose not utilized (Merrill, Jour. Ba,ct.,20, 1930, 235). Fructose, mannitol and trehalose are utilized; sorbitol, arabinose and galactose are not utilized (Gordon, Jour. Bact., 34, 1937, 617). No HoS formed. Optimum temperature 28°C (Kiister), 37 °C (Bynoe). Optimum pH 6.6 to 7.3, range 4.0 to 10.0. Antigenic structure: By agglutination and complement fixation Mycobacterium ranae may be distinguished from Myco- bacterium piscium and Mycobacterium Jriedmannii (Mudd, Proc. Soc. E.xp. Biol, and Med., 23, 1925, 569; Furth, Jour. Immunol., 12, 1926, 286). See Mycobacterium piscium. Pathogenicitj' : Experimentally causes tuberculosis in frogs, lizards, turtles; not pathogenic for rabbits, guinea pigs, rats Or mice. Distinctive characters : See Mycobac- terium piscium. Source : From the liver of a frog. Habitat : In a group of 215 cultures be- longing to the genus, isolated from soils, Gordon (Jour. Bact., 34, 1937, 617) found 65 to sufficiently resemble Mycobacterium ranae to indicate at least a very close relationship. If they prove to be identi- cal, the species is widely distributed. 9. Mycobacteriiun thamnopheos Aron- son. (Jour. Inf. Dis., 44, 1929, 222.) From Thamnophis, a genus of snakes. Tuberculbacillen bei Schlangen, Sib- ley, Arch. f. pathol. Anat. u. Physiol., 116, 1889, 104 {Mycobacterium tropidona- tum (sic) Bergej' et al., Manual, 1st ed., 1923, 376) is probably identical, but the descriptions are too meager to be con- clusive. Acid-fast bacilli described by Gibbes and Shurley (Amer. Jour. Med. Sci., 100, 1890, 145) as the cause of tuberculosis in boas and pythons; by Shattock (Trans. Path. Soc, London, 53, 1902, 430) and by von Hanseneann (Cent. f. Bakt., I Abt., Orig., 34, 1903, 212) as causing tuberculosis in a Python molurus, are possibly identical, but the descriptions do not permit us to draw any conclusions. According to Aronson, similar organisms isolated from patho- logical lesions in boa constrictors and Caluber catenifer differ antigenically from Mycobacterium thamnopheos. Description taken from Aronson {loc. cit.) and Bynoe (Thesis, McGill Uni- versity, Montreal, 1931). Slender rods : 0.5 by 4 to 7 microns, fre- quently slightly curved, beaded and barred forms frequently occur. Non- motile. Acid-fast in cultures of 4 days or older, in younger cultures some or- ganisms are not acid-fast. Not alcohol- fast. Gram-positive. Gelatin stab : Growth occurs along the line of .inoculation. No liquefaction. Agar colonies : 0.5 to 1 mm in diameter, irregular, raised, moist and glistening. Glycerol agar: Spreading, raised, dry, pale pink to buff growth. Glycerol broth : A thin pellicle appears in 5 to 6 days, gradually becomes thicker and falls as a sediment. Dorset's egg medium: Raised, moist, pinkish growth after 10 days, later be- coming salmon-colored. Loeffler's serum : Small, raised, convex, dry growth. Litmus milk: Alkaline. Glycerol potato: Raised, hemispheri- cal, dry and granular growth. Indole not produced. Nitrates : Not reduced by 2 strains, 886 MANUAL OF DETERMINATIVE BACTERIOLOGY reduced by 1 strain (Aronson); slightly reduced (Gordon) ; not reduced (Bynoe). Carbohydrates : Utilizes fructose, man- nitol and trehalose ; fails to utilize arab- inose, sucrose, galactose and sorbitol (Gordon, Jour. Bact., 34, 1937, 617). Temperature relations : Fails to survive 60°C for 1 hour, fails to grow at 47°C (Gordon) ; good growth at 25°C, no growth at 37°C (Aronson); optimum for growth 25°C, range 10° to 35°C (Bynoe). Range of pH : 6.6 to 7.8 (Aronson) ; op- timum 7.3 to 8.0, range 5.0 to 11.0 (Bynoe). Pathogenicity : Experimentally pro- duces generalized tuberculosis in snakes, frogs, lizards and fish but not pathogenic for guinea pigs, rabbits or fowls. Antigenic structure : By agglutination and absorption of agglutinins Mycobacte- rium thamnopheos may be distinguished from Mycobacterium marinum, Myco- bacterium friedmannii and Mycobacterium ranae. See Mycobacterium piscium. Variation: According to Bynoe and Wyckoff (Amer. Rev. Tub., £9, 1934, 389) S and R forms may be distinguished by colony structure and individual cell arrangement . Distinctive characters : See Mycobac- terium piscium. Source : From the lungs and livers of garter snakes (Thamnophis sirtalis). Habitat : Present as a parasite in the garter snake and possibly other cold- blooded vertebrates. 10. Mycobacterium friedmannii Hol- land. (Schildkroten tuberkelbacillus, Friedmann, Cent. f. Bakt., I Abt., Orig., 34, 1903, 647; Bacillus friedmanii (sic) Holland, Jour. Bact., 5, 1920, 218; Myco- bacterium friedmanii Holland, ibid.; Mycobacterium chelonei Bergey et al., Manual, 1st ed., 1923, 376.) Named for Dr. Friedmann, who isolated this or- ganism. Common name-: Turtle bacillus. Description from Friedmann {loc. cit.) and Aronson (Jour. Inf. Dis., 44, 1929, 222). Slender rods: 0.2 to 0.4 by 0.5 to 5 microns. Beaded forms are common. Acid-alcohol -fast in young cultures but in cultures two weeks old generally there are many non-acid-fast rods. Non-mo- tile. Gram-positive. Gelatin stab : White surface growth, scanty growth along the line of stab. No liquefaction. Agar colonies : 1 to 3 mm in diameter, irregularly round, raised, moist, glisten- ing, white. Glycerol agar slants : Thick, spreading growth, at first moist, later granular, 3'^ellowish-white (Friedmann) ; olive-gray (Bynoe); white (Aronson). Glycerol broth : Thick wrinkled pellicle after two to three days growth, later some membranous sediment. Grayish- yellow (Friedmann) ; grayish-white (Bynoe). Dorset's egg medium: Spreading, raised, slightly moist, pale buff. Loeffler's serum : Scant growth, raised, dry, crumb-like. Litmus milk: Slightly alkaline after 10 daj's growth. Glycerol potato: Thick, wrinkled, gray after 2 days growth. Indole not formed. Carbohydrates : Glucose, fructose and arabinose utilized, sucrose slightly uti- lized, galactose and lactose not utilized (Merrill, Jour. Bact., 20, 1930, 235). Arabinose not utilized (Gordon, Jour. Bact., 34, 1937, 617). Optimum temperature 25° to 30°C. Pathogenicity : Experimentally pro- duces tubercles in most species of cold- blooded animals, possibly in guinea pigs but not in other warm-blooded animals. Variation : According to Gildemeister (Cent. f. Bakt., I Abt., Grig., 86, 1921, 513) S and Pi, types may be distinguished on glycerol agar. The S grows as smooth, moist, glistening, convex colonies; the R as flat, dry, spreading colonies. Wykoff (Amer. Rev. Tub., 29, 1934, 289) has shown a difference in the form of cell division and corresponding cell arrange- ment of the two types. FAMILY MYCOBACTERIACEAE \i Distinctive characters : See Mycobac- terium pischnn. Source: From the lungs of turtles in the Berlin aquarium. Habitat : A parasite in turtles and pos- sibly sparingly distributed in soils. Gordon (Jour. Bact., 34, 1937, 617) found 65 out of 215 soil cultures of members of the genus to closely resemble this species. 11. Mycobacteritim spp. (A miscel- laneous group many of which have been incorrectly identified as Mycobacterium leprae Lehmann and Neumann.) Clegg (Phil. Jour. Sci., 4, 1909, 77 and 403), Duval (Jour. Exp. Med., 12, 1910, 649), Duval and Wellman (Jour. Inf. Dis., //, 1912, 116), Currie, Brinckerhoff and Hollmann (Pub. Health Rep., 35, 1910, 1173) and others have described as Myco- bacterium leprae a group of organisms isolated from lepros}^ lesions. Much evidence, summarized by Mclvinley (Med- icine, 13, 1934, 377), points to the conclu- sion that these organisms are not patho- genic and not the causal agent of leprosy. They cannot therefore be included under Mycobacterium leprae as defined above. Thomson (Amer. Rev. Tub., 36, 1932, 162), Gordon (Jour. Bact., 34, 1937, 617), and Gordon and Hagan (Jour. Bact., 36, 1938, 39) recently separated the sapro- phytic members of the genus Mycobac- terium into three main groups and several subgroups. Species names as here de- fined have been added to the key as follows : Group I. Fail to survive 60°C for 1 hour. Grow at 47 °C. a. Utilizes arabinose. Mycobacterium lacticola. b. Unable to utilize arabi- nose. Mycobacterium sp. Group II. Fail to survive 60°C for 1 hour. Do not grow at 47°C. a. Unable to utilize sorbitol. 1. Unable to utilize arab- inose. Mycobacterium, ranac. Mycobacterium thamno- pheos. Mycobacteriuvi sp. 2. Utilize arabinose. Mycobacteritim mar- inum. Mycobacterium sp. h. Utilizes sorbitol. Mycobacterimn spp. c. Unable to utilize most carbohydrates. Mycobacterium fried- mannii. Mycobacterium sp. Group III. Survive 60°C for 1 hour. Grow at 47 °C. a. Utilizes arabinose. Mycobacterium phlei. h. Unable to utilize arabi- nose. Mycobacterium sp. In this study Gordon and Hagan in- cluded many recently isolated soil forms, named saprophytic species, pathogens for cold-blooded animals and 19 cultures, from various collections, which bore the name Mycobacterium leprae. Of these so-called Mycobacterium leprae, six be- long to Group I which corresponds with Mycobacterium lacticola and includes many soil forms, two belong to Group Ila which includes Mycobacterium ranae, Mycobacterium thamnopheos and a num- ber of undefined soil forms, while eleven belong to Group lib. The latter group includes a number of soil cultures but no other defined species. In the several groups to which so-called Mycobacterium leprae strains belong, some appear to be indistinguishable from soil forms, others are distinguished by habit of growth, utilization of carbohy- drates or by pigmentation. 12. Mycobacterimn lacticola Lehmann and Neumann. (Bakt. Diag., 2 Aufl., 3, 888 MANUAL OF DETERMINATIVE BACTERIOLOGY 1899, 409.) From Latin lac, lactis, milk and colo, to dwell ; hence, a milk dweller. From the fact that Lehmann and Neu- mann {loc. cit., 411) refer to the binomial Bacillus friburgensis Korn, it is evident that the species name Jrihurgensis (see Appendix) published the same year (1899) has priority over the species name lacticola. However, since it has never been used with the broad meaning given Mycobacterium lacticola by Lehmann and Neumann in the original description, it is not substituted for the more com- monly used Mycobacterium lacticola in this edition of the Manual. Description from Lehmann and Neu- mann {loc. cit) and Jensen (Proc. Linnean Soc. of New So. Wales, 59, 1934. 19). Slender rods : 0.5 to 0.7 by 2 to 8 microns in young cultures, in older cultures the rods are shorter and frequently coccoid in shape. Curved and irregular forms occur occasionally. Branched forms, if they occur, are ver\- rare. Staining is generally uniform but slight beading occurs occasionally. Strongly acid-fast except organisms from glucose-contain- ing media which are sometimes only faintly acid-fast. Gram-positive. Gelatin colonies: Similar to those on agar. Gelatin stab: Filiform growth in stal). No liquefaction. Agar colonies : Convex, glistening, with entire margins, at first smooth but after 10 to 14 days growth folded or wrinkled. Opaque, at first white, after 2 or 3 days growth becomes yellow. Glucose agar : Similar to agar but more rapid growth and less intensely pig- mented. Glycerol agar slants : Spreading, moist, wrinkled, pale cream-colored to yellow. Nutrient broth: Diffuse growth, later with yellowish pellicle. Litmus milk: Small white granules of growth at the surface, later a dry yellow- ish pellicle. After some weeks' growth the milk becomes alkaline and clear. No coagulation. Dorset's egg medium: As on glycerol agar. Coagulated serum : As qp glycerol agar. Potato: Spreading, raised, wrinkled growth, pale yellow to orange. Long's medium lacking glycerol : No growth. Long's medium with 5 per cent glycerol : Acid formed. (Thomson, Amer. Rev. Tub., 26, 1932, 162.) Indole not formed. Nitrates: Reduced, doubtful (Jensen). Carbohydrates : Glucose, fructose, arab- inose and galactose are utilized ; lactose is not utilized ; sucrose is not utilized by 3 strains, utilized by 1 strain {Mycobac- teriiwi friburgensis) (Merrill, Jour. Bact., 20, 1930, 235). Sorbitol, arabinose, galac- tose, trehalose, mannitol and fructose are utilized ; sucrose is not utilized (Gordon, Jour. Bact., 34, 1937, 617). Optimum temperature 37°C, maximum temperature for growth 52°C, minimum 15° to 18°C. Fails to survive 60°C for 1 hour, grows at 47°C (Gordon, Jour. Bact., 34, 1937, 617; Gordon andHagan, Jour. Bact., 36, 1938, 39). Optimum pH 6.8 to 7.2. Limits for growth 4.5 to 10.0. Distinctive characters: Saprophytic acid-fast organism. Grows rapidly on most media, develops a yellow or orange pigmentation after 3 to 4 days growth. Fails to grow on Long's medium lacking glycerol and produces acid when glycerol is present. Fails to survive 60°C for an hour, grows at temperatures as high as 47°C. Variation : Lehmann and Neumann (Bakt. Diag., 2 Aufi., 2, 1899, 408) and Haag (Cent. f. Bakt., II Abt., 71, 1927, 1) describe three forms : a flat smooth form, a moist, slimy, smooth form and a dry, friable perrugose form. The two former correspond with S and the latter with R types described by Bynoe as characteris- tic of Mycobacterium stercoris, Myco- bacterium berolinensis, Mycobacterium butyricum and Mycobacterium graminis which in turn correspond with S and R types of other members of the genus. Schwabacher (Spec. Rep. Ser. Med. Res. FAMILY MYCOBACTERIACEAE 889 Coun., London, Xo. 182, 1933) finds a difference in the arrangement of the indi- vidual cells of the S and R types. Source : From butter, plant dust, cow manure . Habitat : Gordon (Jour. Bact., 34, 1937, 617) found 1 culture isolated from nasal exudate, 1 from bovine lymph gland and 94 isolated from soil, out of a group of 215 soil cultures belonging to the genus, to be either identical with or very closely related to this species. If these strains are valid members of the species, Myco- bacterium lacticola is widely distributed in soil, dust, dairy products, etc. 13. Mycobacterium phlei Lehmann and Neumann. (Timotheebaciilus or Gras- bacillus I, Moeller, Iherapeutischen Monatsheften, 12, 1898, 607; IMoeller, Deut. med. Wclmschr., 24, 1898, 376; Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 411; Mycobacterium moelleri Chester, Manual Determ. Bact., 1901, 358; Sclerothrix phlei Vuillemin, Encyclopedie Mycologique, Paris, 2, 1931, 160.) From :^I. L. Phleum, a genus of grasses. Description from Moeller (loc. cit.) and Jensen (Proc. Linnean Soc. New So. Wales, 59, 1934, 32). Slender rods : 0.2to 0.5 by 1 to 4 microns, sometimes club-shaped, frequently beaded, rarelj^ branched. Strongly acid- fast and acid -alcohol -fast in cultures older than 2 to 3 days, in younger cultures there are generally many non-acid-fast cells. Non-motile. Gram-positive. Gelatin colonies : Small, 0.5 to 1 mm in diameter; irregular, raised, moist and glistening, finely granular, orange. Gelatin stab : Filiform, opaque, orange. No liquefaction. Agar colonies : Similar to gelatin colo- nies, yellow to orange. Agar slant : Spreading, raised, dry with roughened granular surface, yellow to orange. Broth: Turbid, with yellow pellicle. Dorset's egg medium: Spreading, raised, dry, orange. Loeffier's serum: Similar to Dorset's egg medium, creamy to j-ellow. Glj'cerol broth : Thin transparent pelli- cle, later becoming thickened, rough, wrinkled and yellow to pink, still later a flaky sediment. Litmus milk : Yellow flocculi on sur- face, slowly becomes alkaline. No co- agulation. Potato: Thick, dry, j'cllow, adherent growth. Long's medium lacking glycerol : Abun- dant growth. Long's medium with 5 per cent glycerol : No acid formed (Thomson, Amer. Rev. Tub., 26, 1932, 162). Nitrites are produced from nitrates. Indole not formed. Carbohydrates : Glucose, fructose, arab- inose, trehalose, mannitol and galactose are utilized; sucrose and lactose are not utilized (Merrill, Jour. Bact., 20, 1930, 235; Gordon, Jour. Bact., 34, 1937, 617). Temperature relations : Survives 60°C for 1 hour, grows at 47°C (Thomson, Amer. Jour. Tub., 26, 1932, 162); opti- mum for growth 37°C, range 20° to 58°C (Bynoe). Optimum pH 6.8 to 7.3; range 5.5 to 10.0. Pathogenicitj- : The injection of large numbers of organisms into guinea pigs results in a local abscess of a few weeks' duration, occasionally small abscesses de- velop in the regional lymph glands or the visceral organs. According to Mayer (Cent. f. Bakt., I Abt., 26, 1899, 331) and others, the injection of the organisms along with butter or other fat increases the pathological reaction. Variation: Haag (Cent. f. Bakt., II Abt., 71, 1927, 1) and Bynoe (Thesis, McGill University, Montreal, 1931) find two or three colony types : an R form which fits into the description of the species given above and an S type which grows as a perfectlj' smooth, raised, moist, glistening colony with an entire margin. Cooper (Jour. Inf. Dis., 54, 1934, 236) distinguished pigmented and non-pigmented types. 890 MANUAL OF DETERMINATIVE BACTERIOLOGY Distinctive characters: Saprophyti acid-fast organism, grows rapidly on most media. Shows yellow pigmentation as soon as growth is visible. Grows well on Long's medium lacking glycerol and fails to produce acid when glycerol is present. Survives 60°C for 1 hour and grows at 47°C. Source: Originally isolated from hay and grass. Frequently found in soil, dust and other sources. Out of 215 cultures of the genus recovered from soils by Gordon (Jour. Bact., 34, 1937, 617) Mycobacterium phlei was isolated on 22 occasions. The same author re- ports 3 cultures of a closely related if not identical organism recovered from bovine lymph glands, 1 recovered from bovine skin and 1 recovered from a hen's spleen. Habitat: Widely distributed in soils, dust, hay, etc. Appendix I : The following saprophytic species have been placed in this genus. Their relationships are not clear. Some are related to or possibly identical with Mycobacterium lacticola. Mycobacterium album Sohngen. (Cent. f. Bakt., II Abt., 37, 1913, 599.) From garden earth. Mycobacterium bekkerii Bekker. (An- tonie van Leeuwenhoek, 9, 1943, 81 ; abst. in Cent. f. Bakt., I Abt., Orig., 149, 1944, 500.) From urine. Mycobacterium berolinense Bergey et al . (Tuberkelahnlichen Bacillen, Rabino- witsch, Ztschr. f. Hyg., 26, 1897, 90; Mycobacterium lacticola /3 perrugosum Lehmann and Neumann, Bakt. Diag., 2 Aufl., 3, 1899, 410; Mycobacterium lacti- cola perrugosum Haag, Cent. f. Bakt., II Abt., 71, 1927, 3 ; Bergey et al.. Manual, 1st ed., 1923, 377.) From butter. Mycobacterium butyri Chester. (Man. Determ. Bact., 1901, 357.) This name includes both the Tuberkelahnlichen Bacillen of Rabinowitsch and the Butter Bacillus of Petri. From butter. Mycobacterium butyricum Bergey et al. (Butter Bacillus, Petri, Arb. kaiserl. cGesundheitsamte, 14, 1898, 1 ; Bergey et al., Manual, 1st ed., 1923, 377.) From butter. Mycobacterium cholesterolicum Tak. (Antonie van Leeuwenhoek, 8, 1942, 39.) From garden soil. Mycobacterium friburgensis (Korn) Chester. (Bacillus friburgensis Korn, Cent. f. Bakt., I Abt., 25, 1899, 532; Mycobacterium lacticola y friburgensis Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899,411 ; Chester, Man. Determ. Bact., 1901, 359.) From butter. Mycobacterium graminis Chester. (Grasbacillus II, Moeller, Cent f. Bakt., I Abt., 25, 1899, 369; Mycobacterium lac- ticola a planum Lehmann and Neumann, Bakt. Diag., 2 Aufl., 2, 1899, 408; Chester, Man. Determ. Bact., 1901, 358; Myco- bacterium lacticola planum Haag, Cent, f. Bakt., II Abt., 71, 1927, 3.) From hay dust. Mycobacterium hyalinum Sohngen. (Cent. f. Bakt., IFAbt., 37, 1913, 599.) From garden earth. Mycobacterium luteum Sohngen ijioc. cit.). From garden earth. Mycobacterium muris Simmons. (Jour. Inf. Dis., 4^, 1927, 13.) From the feces of gray mice. Mycobacterium phlei perrugosum Haag. (Cent. f. Bakt., II Abt., 71, 1927, 6.) From soils and manure. Mycobacterium phlei planum Haag (loc. cit.). From soils. Mycobacterium ranicola /and II Haag {loc. cit.). From frogs. Mycobacterium rubrum Sohngen {loc. cit.). From garden earth. Mycobacterium smegmalis (Trevisan) Chester. (Smegma bacillus, Alvarez and Tavel, Arch. Phys. norm, et path., 6, 1SS5, 303; Bacillus smegmatis Trevisan, I generi e le specie delle Batteriacee, 1889, 14; Bacterium smegmatis Migula, Syst. d. Bakt., 2, 1900, 497; Chester, Man. Determ. Bact., 1901, 357.) From smegma. Weber (Arb. kaiserl. Gesund- heitsamte, 19, 1902, 251) finds Mycobac- terium smegmatis acid- but not alcohol- fast in contrast to the mammalian FAMILY MYCOBACTERIACEAE 891 tubercle bacilli which are both acid- and alcohol-fast. Later observers (Bynoe, Thesis, McGill University, Montreal, 1931) have not found this a valid dis- tinction. Mycohacteriuni smegmatis var. muris Galli-Valerio. (Cent. f. Bakt., I Abt., Orig., 75, 1915, 49.) From the preputial glands of the black rat {Mus rattus) . Mycobacterium stercoris Bergey et al. (Mist Bacillus, Moeller, Berlin, thier- arztl. Wochnschr., 1898, 100; Mycobac- terium stercusis (sic) Bergey et al., Manual, 1st ed., 1923, 378; Bergey et al., Manual, 4th ed., 1934, 542.) From manure . Mycobacterium testudims Friedmann and Piorkowski. (See Haag, Cent. f. Bakt., II Abt., 71, 1927, 5; apparently the same as Mycobacterium testudo, loc. cit., 10.) This is probably Mycobacte- rium friedmannii. From turtles. Appendix II: Krassilnikov (Mikro- biol., 7, 1938, 335; and Ray Fungi and Related Organisms, Izd. Acad. Nauk. Moskow, 1938, 121-130) describes a genus Mycococcus distinct from Hans- girg's (Osterr. Bot. Ztschr., S8, 1888, 266) family Mycococcaceae (which is related to the fungi) and distinct from Mycococcus Bokor (Arch. f. Mikrobiol., 1, 1930, 1). Mycococcus Krassilnikov includes species that produce coccus-like cells, genetically related to the species included in Mycobacterium; reproduction is by fission or budding in different directions, often forming short, irregular chains with side branches; in old cultures, the vege- tative cells change into resting cells, the latter germinating in a manner similar to the spores of actinomycetes. Seven species are listed, with incomplete de- scriptions. Mycococcus ruber, M. capsu- latus, M. luteus, M. citreus and M. albus are described in Krassilnikov's original paper. One of these {Mycococcus luteus) is dropped in his later monograph while descriptions of two new species are added (Mycococcus tetragenus and M. mucosus). 892 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY II. ACTINOMYCETACEAE BUCHANAN.* (Jour. Bact., 3, 1918, 403.) Mycelium is non-septate during the early stages of growth but later may become septate and break up into short segments, rod -shaped or spherical in shape, or the mycelium may remain non-septate and produce spores on aerial hyphae. The or- ganisms in culture media are either colorless or produce various pigments. Some species are partially acid-fast. This family is distinguished from the previous one by the formation of a true mycelium. As compared with the next family, it is charac- terized by the manner of spore formation. Key to the genera of family Actinomycetaceae. I. Obligate aerobic. The colonies are bacteria-like in nature, smooth, rough or folded, of a soft to a dough-like consistency, sometimes compact and leathery in young stages. Most forms do not produce any aerial mycelium; a few pro- duce a limited mycelium, the branches of which also break up into oidiospores or segmentation spores. Some species are partially acid-fast. Genus I. Nocardia, p. 892. II. Anaerobic or microaerophilic, parasitic; non-acid -fast, non-proteolytic and non- diastatic. Genus II. Actinomyces, p. 925. Genus I. Nocardia Trevisan. (Trevisan, I generi e le specie delle Batteriacee, 1889, 9; Actinomyces Gasperini, Cent. f. Bakt., 15, 1894, 684 and Atti dell' XI congresso med. internaz. Roma, 6, 1895, 82; not Actinomyces Harz, Jahresber. d. Miinch. Thierarzneischule for 1877- 1878, 1879, 125; Actitiobacter i2im Haas , Cent. f. Bakt., I Abt., Orig., 40, 1906, 180; Sam- pietro, Ann. d'Igiene, Roma, 18, 1908, 331; Actinococcus Beijerinck, Folio Microbiol., Delft, 3, 1914, 196; not Actinococcus Kiitzing, Species Algarum, 1849; Brevistrepto- thrix Lignieres, Ann. Parasit. hum. et comp., 2, 1924, 1 ; Asteroides Puntoni and Leo- nardi. Boll, e Atti d. R. Accad. Med., 67, 1935, 90; Proactinomyces Jensen, Proc. Linn. Soc. New So. Wales, 56, 1931, 345.) Named for Prof. E. Nocard who first described the type species. Slender filaments or rods, frequently swollen and occasionally branched, forming a mycelium which after reaching a certain size assumes the appearance of bacterium- like growths. Shorter rods and coccoid forms are found in older cultures. Conidia not formed. Stain readily, occasionally showing a slight degree of acid-fastness. Non -motile. No endospores. Aerobic. Gram-positive. The colonies are similar in gross appearance to those of the genus Mycobacterium. Paraffin, phenol and m-cresol are frequently utilized as a source of energy. In their earlj- stages of growth on culture media (liquid or solid), the structure of nocardias is similar to that of actinomyces in that they form a tj'pical mycelium hyphae branch abundantly, the branching being true. The diameters of the hyphae vary between 0.5 and 1 micron, usually 0.7 to 0.8 micron, according to the species. The mycelium is not septate. However, the further development of nocardias differs sharply from that of actinomyces : the filaments soon form transverse walls and the whole mycelium breaks up into regularly cylindrical short cells, then into coccoid * Completely revised by Prof. S. A. Waksman, New Jersey E.xperiment Station, New Brunswick, New Jersey and Prof. A. T. Henrici, University of Minnesota, Minneapolis, Minnesota, May, 1943. FAMILY ACTINOMYCETACEAE 893 cells. On fresh culture media, the coccoid cells germinate into mycelia. The whole cycle in the development of nocardias continues for 2 to 7 days. Most frequently the coccoid cells are formed on the third to fifth day, but in certain species (e.g., Nocardia rubra) they can be found on the second day. Numerous chlamydospores may be found in older cultures of nocardias. They are formed in the same way as the chlamydospores in true fungi: the plasma inside the filaments of the nn'celium condenses into elongated portions. In older cultures of nocardias many coccoid cells are changed into resistant cells. The latter are larger than the vegetative coccoid cells ; the plasma of these cells is thicker than the plasma of vegetative cells ; on fresh media they germinate like the spores of actinomyces ; they form 2 to 3 germ tubes. Besides the cells mentioned, numerous involution forms can often be found in older cultures of nocardias ; the cells are thin, regularlj^ cylindrical or coccoid, are often transformed into a series of spherical or elliptical ampules and a club-like form (2 to 3 microns and more) . The multiplication of nocardias proceeds by fission and budding; occasionally they form special spores. Budding occurs often. The buds are formed on the lateral surface of the cells ; when they have reached a certain size, they fall off and develop into rod-shaped cells or filaments. The spores are formed by the breaking up of the cell plasm into separate portions usualh' forming 3 to 5 spores ; every portion becomes rounded, covered with a membrane and is transformed into a spore; the membrane of the mother cell dissolves and disappears. The spores germinate in the same way as those of actinomyces. They form germ tubes which develop into a mycelium. The colonies of nocardias have a paste-like or mealy consistency and can easily be taken up with a platinum loop ; they spread on glass and occasionally render the broth turbid. The surface colonies are smooth, folded or wrinkled. Typical nocardias never form an aerial mycelium, but there are cultures whose colonies are covered with a thin coating of short aerial hj^phae which break up into cylindrical oidiospores. Many species of nocardias form pigments; their colonies are of a blue, violet, red, yellow or green color; more often the cultures are colorless. The color of the cul- ture serves as a stable character. . Krassilnikov (Raj- fungi and related organisms, Izd. Acad. Nauk, U.S.S.R., Moscow, 1938) divides the genus into two groups: 1. Well developed aerial mycelium; sub- strate mycelium seldom produces cross-walls ; the threads break up into long, thread- like rods; branches of the aerial mycelium produce segmentation spores and oidio- spores; the latter are cjdindrical with sharp ends; no spirals or fruiting branches. This group is the same as group B of Jensen (loc. cit.). 2. Typical forms; mycelium develops only at early stages of growth, then breaks up into rod-shaped and coccoid bodies ; smooth and rough colonies, dough-like consistency ; never form an aerial myce- lium; similar to bacterial colonies ; aerial mycelium may form around colonies. This genus can also be divided, on the basis of acid-fastness, into two groups: Group 1. Partially acid-fast organisms, which are non-proteolytic, non-diastatic and utilize paraffin; usually j^ellow, pink, or orange-red in color. Group 2. Non-acid-fast or- ganisms, which are diastatic, largely proteolytic and do not utilize paraffin; yellow, orange to black in color. The type species is Nocardia farcinica Trevisan. Keij to the species of genus Nocardia. I. Partially acid-fast* organisms with strongly refractive cells; non-proteolytic and generally non-diastatic; constantly capable of utilizing paraffin. * Acid-fastness is not marked in cultures, is apparent in infected tissues, pro- nounced in sputum or other e.xudates 894 MANUAL OF DETERMINATIVE BACTERIOLOGY A. Initial mycelium well developed, richly branching, dividing into rods and generally into cocci. 1. Vegetative mycelium soft, without macroscopically visible aerial mycelium, a. Vegetative mycelium yellow, orange or red. b. Pathogenic. c. Vegetative mycelium white, buff, or pale yellow. 1. Nocardia farcinica. cc. Vegetative mycelium yellow to red. 2. Nocardia asteroides. bb. Not pathogenic. 3. Nocardia polychromogenes . aa. Vegetative mycelium white to pink. b. Gelatin not liquefied. c. Growth on nutrient agar opaque, cream-colored; coccoid forms in broth. 4. Nocardia opaca. cc. Growth on nutrient agar watery, no coccoid forms in broth. 5. Nocardia eryihropolis. ccc. Growth on nutrient agar pink. d. White aerial mycelium on milk. 6. Nocardia leishmanii . ■ dd. Pink pellicle on milk. 7. Nocardia caprae. ddd. Yellow pellicle on milk. 8. Nocardia preloriana. bb. Gelatin liquefied. 9. Nocardia pulmonalis. 2. Vegetative mycelium hard, yellow, with white aerial mycelium ; hyphae divide into chains of acid-fast cocci. 10. Nocardia paraffinae. B. Initial mycelium very short, rapidly dividing into rods and cocci. 1. Slowly growing organisms; cells 0.5 to 0.7 micron in diameter. 11. Nocardia minima. 2. Rapidly growing organisms; cells 1.0 to 1.2 microns in diameter. a. Growth pink. b. Cystites (swollen cells) not formed, c. No indigotin from indole. 12. Nocardia corallina. cc. Indigotin from indole. 13. Nocardia globerula. bb. Cystites formed. 14. Nocardia salmonicolor . aa. Growth coral red. 15. Nocardia rubroperlincia. aaa. Growth dark red. 16. Nocardia rubra. aaaa. Growth white. b. No aerial mycelium. 17. Nocardia coeliaca. bb. Aerial mycelium. 18. Nocardia transvalensis . FAMILY ACTINOMYCETACEAE 895 II. Non-acid-fast organisms with weakly refractive cells; no distinct formation of cocci. Constantly diastatic. A. Not proteolytic. 1. Growth on agar pale cream. 19. Nocardia mesenterica. 2. Growth on agar yellow. 20. Nocardia flava. 3. Growth on agar green. 21. Nocardia viridis. 4. Growth on agar yellow-green. 22. Nocardia citrea. 5. Growth on agar pink to crimson. 23. Nocardia madurae. 6. Growth consistency soft, sparse aerial mycelium. 24. Nocardia lutea. 7. Growth consistency medium, good aerial mycelium. 25. Nocardia blackwellii. 8. Grood action on milk. Growth consistency firm, liberal, aerial myce- lium. 26. Nocardia cuniculi. 9. Deep brown pigment on protein media. 27. Nocardia rangoonensis . 10. Light brown pigment on protein media. 28. Nocardia caviae. B. Proteolytic. 1. Growth on nutrient agar with rapid formation of unbranched diphthe- roid-like rods ; no typical cystites ; broth turbid. 29. Nocardia actinomorpha. 2. Growth on nutrient agar with extensive mycelia; simple unbranched rods not formed; cystites present. Broth clear. 30. Nocardia flavescens. 3. Colonies orange-yellow to orange-red, which may change to black. 31. Nocardia maculata. 4. Light brown pigment on protein media. 32. Nocardia rhodnii. 5. Green to greenish -brown pigment on protein media. 33. Nocardia gardneri. 1. Nocardia farcinica Trevisan. (Ba- perini, ibid.; Cladothrix farcinica Mace, cille du farcin, Nocard, Ann. Inst. Past., Traite de Bacteriologie, 3rd ed., 1894, ;2, 1888, 293; Trevisan, I generi e le specie 1047; Streptothrix farcini bovis Kitt, delle Batteriacee, Milan, 1889, 9; (Sirepio- Bakterienkunde und pathologische trix farcinica Rossi Doria, Ann. d. 1st. Mikroskopie, Vienna, 3 Aufl., 1899, 511; d'Igi. Sper. Univ. di Roma, 1, 1891, 424; Bacteriurn nocardi Migula, Syst. d, Actinomyces farcinicus Gasperini, Ann. Bakt., 2, 1900, 345; Streptothrix nocardii 1st. d'Igiene, Roma, ;2, 1892, 222; Oospora Foulerton, .Jour. Compt. Path, and farcinica Sauvageau and Radais, Ann. Therap., 14. 1901, 51; Discomyces far- Inst. Past., 6y 1892, 248; Actinomyces cinicus Gedoelst, Les champignons feom/amnicus Gasperini, Cent. f. Bakt., parasites de I'homme et des animaux 15, 1894, 684; Bacillus farcinicus Gas- domestiques, Brussels, 1902, 167; Ac- 89G MANUAL OF DETERMINATIVE BACTERIOLOGY tinomyces nocardii Buchanan, Veterinary Bacteriology, Philadelphia, 1911, 378; Nocardia albida Chalmers and Christ o- pherson, Ann. Trop. Med. and Parasit., 10, 1916, 271, according to Dodge, Medi- cal Mycology, St. Louis, 1935, 746.) From M. L. farcinicus, of farcy. Filaments 0.25 micron in thickness, branched. Markedly acid-fast. Gelatin colonies : Small, circular, trans- parent, glistening. Gelatin stab : No liquefaction. Agar colonies : Yellowish-white, irregu- lar, refractive, filamentous. Agar slant : Grayish to yellowish-white growth, surface roughened. Broth: Clear, with granular sediment, often with gray pellicle. Litmus milk: Unchanged. Potato: Abundant, dull, crumpled, whitish-yellow growth. Nitrites not produced from nitrates. No soluble pigment formed. Proteolytic action absent. Starch not hydrolyzed. Aerobic, facultative. Optimum temperature 37°C. Source: From cases of bovine farcy. Habitat : Associated with a disease in cattle, resembling chronic tuberculosis. Transmissible to guinea pigs, cattle and sheep, but not to rabbits, dogs, horses or monkeys . 2. Nocardia asteroides (Eppingerj Blanchard. {Cladothrix asteroides Ep- pinger, Beitr. z. path. Anat., 9, 1891, 287; Slreptotriz (sic) eppingerii Rossi Doria, Ann. Inst. d'Ig. sper. d. Univ. Roma, 1, 1891, 423; Streptotrix (sic) asteroides Gasperini, Ann. Inst. d'Ig. sper. d. LTniv. Roma, :2, 1892, 183; Oospora asteroides Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 252; Actinomyces asteroides Gasperini, Cent. f. Bakt., 15, 1894, 684; Blanchard, in Bouchard, Traite Path. G6n., 2, 1895, 811; Discomyces asteroides Gedoelst, Champ. Paras. Homme et Anim., 1902, 173, Actinomyces eppingeri Namvslowski, Cent. f. Bakt., I Abt.. Orig., 62, 1912, 566; Asteroides asteroides Puntoni and Leonardi, Boll, e Atti d. R. Accad. Med. di Roma, 61, 1935, 92; Mycobacterium asteroides, quoted from Puntoni and Leonardi, idem; Proacti- nomyces asteroides Baldacci, Soc. Internat. di Microb., Boll. d. Sez. Ital., 9, 1937, 141.) From Greek aster, star and cidos, shape. Probable synonyms : Streptotrix auran- tiaca Rossi Doria, loc. cit., 417 (Actino- myces aurantiacus Gasperini, loc. cit., 1892, 222; Oospora aurantiaca Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 388; Cladothrix aurantiaca Mace, Traite Pratique de Bact., 4th ed., 1901, 1096; Nocardia aurantiaca Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268) and Streptothrix freeri Musgrave and Clegg, Philippine Jour. Sci., Med. Sciences, 2, 1907, 477 (Discomyces freeri Brumpt, Precis de Parasitol., Paris, 1st ed., 1910, 858, Nocardia freeri Pinoy, Bull. Inst. Past., 11, 1913, 936; Oospora freeri Sartory, Champ. Paras. Homme et Anim., 1923, 785; Actinomyces freeri Bergey et al., Manual, 1st ed., 1923, 346). According to Chalmers and Christopherson (loc. cit.) another synonym of this organism is Streptothrix hominis Sabrazes and Riviere, Le Semaine M^decine, 1895, no. 44. Straight, fine mycelium, 0.2 micron in thickness, which breaks up into small, coccoid conidia. Acid-fast. Gelatin stab : Yellowish surface growth. No growth in stab. No liquefaction. Synthetic agar : Thin, spreading, orange growth. No aerial mycelium. Starch agar: Restricted, scant, orange growt h . Plain agar: Much folded, light yellow- growth, becoming deep yellow to yellow- ish-red. Glucose broth : Thin, yellowish pellicle. Litmus milk : Orange -colored ring. No coagulation. No peptonization. Potato: Growth much wrinkled, whit- ish becoming vellow to almost brick-red. FAMILY ACTINOMYCETACEAE 897 Nitrites produced from nitrates. No soluble pigment formed. Proteolytic action doubtful. Starch not hydrolyzed. Transmissible to rabbits and guinea pigs but not to mice. Aerobic. Optimum temperature 37°C. Source : From a cerebral abscess in man . Habitat : Also found in conditions re- sembling pulmonary tuberculosis. A number of strains of acid-fast ac- tinomycetes isolated from human lesions have deviated in certain particulars from the description of Nocardia asteroides, but not sufficiently to warrant separation as species. The following varieties are described by Baldacci (Mycopathologia, 1, 1938, 68): Nocardia asteroides var. crateriformis (Baldacci) comb. nov. (.Proactinomyces asteroides var. crateriformis Baldacci, loc. cit.) Less tendency to fragmentation of mycelium. Complete lack of aerial mycelium. Growing as discrete colonies, disk- or crater-shaped. Nocardia asteroides var. decolor (Baldacci) comb. nov. (Proactinomyces asteroides var. decolor Baldacci, loc. cit.) Greater tendency to produce white aerial mycelium; vegetative mycelium colorless . Nocardia asteroides var. gypsoides (Baldacci) comb. nov. {Actinomyces gypsoides Henrici and Gardner, Jour. Inf. Dis., 28, 1921, 248; Discomyces gypsoides Brumpt, Precis de Parasitol., Paris. 3rd ed., 1922, 980; Oospora gypsoides Sartory, Champ. Paras. Homme et Anim., 1923, 802; Proactinoinyces as- teroides var. gypsoides Baldacci, loc. cit.) White aerial mycelium; darkening of peptone media. 3. Nocardia polychromogenes (Vallee) comb. nov. {Streptothrix polychromogenes Vallee, Ann. Inst. Past., 17, 1903, 288; Streptothrix pluricromogena Caminiti, Cent. f. Bakt., I Abt., Orig., U, 1907, 198; Actinomyces polychromogenes Lieske, Morphol. u. Biol. d. Strahlenpilze, Leipzig, 1921, 32; Proactinomyces poly- chromogenes Jensen, Proc. Linnean Soc. New So. Wales, 56, 1931, 79 and 363; Oospora polychromogenes Sartorj-^, quoted from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51; Acti- nomyces plurichromogenus Dodge, Medi- cal Mycology, St. Louis, 1935, 737.) From Greek, producing many coloi's. Description from Jensen (loc. cit.). Long wavy filaments : 0.4 to 0.5 by 70 to 100 microns, extensively branched but without septa. Older cultures consist entirely of rods 4 to 10 microns, fre- quently in V, Y, or smaller forms. Still older cultures consist of shorter rods and coccoid forms. Gram-positive, fre- quently showing bands and granules. Gelatin stab : Thin yellowish growth along the stab with thin radiating fila- ments. Surface growth flat, wrinkled, red. No liquefaction. Nutrient agar: Scant, orange-red growth. Glucose agar: After 3 to 4 days raised, flat, glistening, rose -colored growth. After 1 to 3 weeks becoming folded and coral -red. Glucose broth : After 3 to 4 days tur- bid; after 2 to 3 weeks an orange flaky sediment. No surface growth. Milk : Growth starts as small orange- colored surface granules. After 1 to 2 weeks a thick, soft, orange -colored sedi- ment forms. Optimum temperature 22° to 25°C. Distinctive characters : Differs from Nocardia corallina in the formation of very long filaments and in filiform growth in gelatin stabs. Source: From the blood of a horse; from soil in France and Australia. Habitat: Soil. 4. Nocardia opaca (den Dooren de Jong) comb. nov. {Mycobacterium opacum den Dooren de Jong, Cent. f. Bakt., II Abt., 71, 1927, 216; Mycobacterium crys- tallophagum Gray and Thornton, Cent. 898 MANUAL OF DFiTERMIXATIVE BACTERIOLOGY f. Bakt., II Abt., 73, 1928, 86; Actino- myces crystallophagus Bergey et al., Manual, 3rd ed., 1930, 473; Proactino- myces opac us Jensen, Proc. Linnean Soc. New So. Wales, 57, 1932, 369; Proaciino- myces crystallophagus Reed, in Manual, 5th ed., 1939, 834.) From Latin opacus, shady, darkened. Description from Gray and Thornton {loc. cit.), Bynoe (Thesis, McGill Uni- versity, Montreal, 1931), and Jensen {loc. cit.). Long, curved, irregular and branching filaments or rods: 0.8 to 1.0 by 2 to 16 microns, or occasionally longer. Few chains or clumps are formed. In older cultures shorter rods or cocci are gener- ally formed. Readily stained. Not acid-fast. Gram-positive. Gelatin colonies : Round, convex, whit- ish, smooth, shining, edges slightly ar- borescent. Deep colonies: Burrs, with slightly irregular processes. Gelatin stab : Convex, whitish, smooth, resinous, filiform, erose. Broth : Turbid with broken white scum, or clear with granular suspension. Dorset's egg medium: Spreading, smooth, moist, salmon-colored growth. Loeffler's medium: Scanty growth, smooth, moist, light buff-colored. Glycerol potato : Dry, rough, crumpled, pink to buff -colored growth. Litmus milk : Grayish pellicle ; slightly alkaline. Nitrites are produced from nitrates. No acid from sucrose, lactose, maltose or glucose. Phenol and naphthalene are utilized as sources of energy. Optimum pH 6.8 to 7.3. Optimum temperature 30°C. Distinctive characters : Differs from Nocardia corallina and Nocardia poly- chromogenes in that the cells are much longer than those of the former and much shorter than those of the latter. Grows in smooth convex surface colonies and burr-like deep colonies. Source : Twenty-four strains isolated from soils in Great Britain. Habitat : Probably sparingly distrib- uted in soils. 5. Nocardia erythropolis (Gray and Thornton) comb. nov. {Mycobacterium erythropolis Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 87; Actino- myces erythropolis Bergey et al., Manual, 3rd ed., 1930, 472; Proactinomyces erythropolis Jensen, Proc. Linnean Soc. of New So. Wales, 57, 1932, 371.) From Greek erythrus, red and polis, city. Description from Gray and Thornton {loc. cit.) and from Bynoe (Thesis, McGill University, Montreal, 1931). Long uneven-sided rods and filaments, curved and branching up to 11 microns long by 0.8 micron. Coccoid forms not formed. Stains readily. Not acid-fast. Gram-positive. Gelatin colonies : After 12 days, round, flat, white, shining; edge entire. Deep colonies: Round, smooth. Gelatin stab: After 8 to 14 days, con- vex, white, smooth, shining, radiate from center, borders cleft. Line of puncture filiform, erose. Agar colonies : Round, 2 to 3 mm in diameter, convex, watery -white ; edge entire. Deep colonies : Lens-shaped. Agar slant : Filiform, flat, watery growth; edge undulate. Broth: Growth slight, turbid. Dorset's egg medium: After 2 weeks, raised, moist, finely granular, irregular margin, flesh-colored. Loeffler's medium : After 7 days growth as on Dorset's egg medium, but pink. Glycerol potato: After 7 days, flat, dry, rough, orange -colored. Litmus milk: Pale pink pellicle. Nitrites not produced from nitrates. No acid from glucose, lactose, sucrose or glycerol. Phenol is utilized. Optimum pH 6.8 to 8.0. Optimum temperature 25°C. FAMILY ACTINOMYCETACBAE 899 Distinctive characters : Differs from Nocardia coeliaca and Nocardia actino- morpha in the filiform growth and absence of liquefaction of gelatin. Long rods and filaments . Source : Six strains isolated from soils in Great Britain. Habitat: Presumablj' soil. 6. Nocardia leishmanii Chalmers and Christopherson. (A new acid-fast streptothri.x, Birt and Leishman, Jour. Hyg., ^, 1902, 120; Chalmers and Christo- pherson, Ann. Trop. Med. and Parasit., 10, 1916, 255; Discomyces leishmani Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 984; Actinomyces leishmani Sartory and Bailly, Mycoses pul- monaires, 1923, 253.) Named for Leish- man, one of the original isolators. Description from Erikson {loc. cit., p. 27). Initial cells frequently swollen, large and irregular, aggregated in short chains and then branching out into regular narrow filaments ; at margin of colony on synthetic glycerol agar may be seen comparatively long thick segments with accompanying fringe of normal hyphae ; later entire colonies asteroid in appear- ance, very fine and close angular branch- ing, with aerial hyphae situated singly; aerial mycelium generally abundant with irregularly cylindrical conidia. Slightly acid-fast. The latter property must have been attenuated during artificial cultivation, for the organism is reported as markedly acid-fast by the original isolators. Gelatin : Small pink colonies in depths of stab. No liquefaction. Glucose agar: Rounded elevated colo- nies with paler frosting of aerial mycelium ; grow^th becoming piled up, aerial myce- lium sparse. Glycerol agar: Small round pink colo- nies, tending to be umbilicated and piled up, stiff w^hite aerial spikes. Coon's agar : Small round colorless colonies, stiff white aerial spikes; later a pink tinge. Potato agar : Minute colorless round colonies, small raised patches of white aerial mycelium. Dorset's egg medium: Colorless con- fluent growth studded with little wart- like projections bearing stiff aerial spikes ; growi;h becomes pinkish with a white aerial mycelium; later, growth drab gray, medium discolored. Serum agar: Minute round colorless colonies with pinkish tinge in confluent raised patch. laspissated serum : Small round pale pink colonies, umbilicated and raised up. Broth: Liberal growth, white floccu- lent colonies; later pink surface colonies. Synthetic sucrose solution : Colorless flocculent sediment, thin colorless pellicle. Milk: Surface growth, white aerial mycelium, solid coagulum; later partly peptonized with pink aerial mycelium. Litmus milk: Pink surface growth, aerial mycelium, milky opaque after 40 days. Carrot plug: Small irregularly round raised colonies, colorless, covered vath stiff aerial spikes ; later buff-colored, con- voluted and ribbed growth with small patches of white aerial mycelium; aerial mycelium pink in two months. Source : From fatal case of lung disease and pericarditis in man. Habitat : Human infections so far as laiown . 7. Nocardia caprae (Silberschmidt) comb. nov. (Streptothrix caprae Silber- schmidt, Ann. Inst. Past., 13, 1899, 841; Cladothrix caprae Mace, Traite Pratique de Bact., 4th ed., 1901, 1094; Discomijces caprae Gedoelst, Champ. Paras. Homme et Anim., 1902, 174; Oospora caprae Sartory, Champ. Paras. Homme et Anim., 1923, 813; Actinomyces caprae Ford, Textb. of Bact., 1927, 205.) From Latin capra, goat. 930 MANUAL OF DETERMINATIVE BACTERIOLOGY Description from Erikson (Med. Res. Council Spec. Rept. Ser. 203, 1935, 26). Initial cells only slightly enlarged; early development of aerial hyphae, while substratum threads are still short ; frequent slipping of branches ; aerial mycelium abundant on all media with tendency to form coherent spikes; my- celium not very polymorphous, but occasional thicker segments appear. Slightly acid -fast. Gelatin: Extensive dull growth with small raised patches of pink aerial myce- lium; later ribbon-like, depressed. No liquefaction. Glucose agar ; Irregular bright pink growth tending to be heaped up; later abundant masses frosted over with thin white aerial mycelium. Glycerol agar : Abundant growth, small round pink colonies, partly covered with white aerial mycelium. Potato agar: Extensive thin growth, pink in raised patches, covered by white aerial mycelium; later aerial mycelium also becomes pink. Starch agar : Minute colorless colonies covered by white aerial mycelium. Blood agar : Minute round colorless colonies aggregated in broad pink zones, paler aerial mycelium. No hemolysis. Dorset's egg medium: Few colorless colonies, some pink, white aerial myce- lium; later, growth becoming dull pink, irregular, with scant white aerial myce- lium. Ca-agar: Minute colorless colonies, white aerial mycelium; later a pinkish tinge. Serum agar : Small round pink colonies frosted over with thin white aerial my- celium. Inspissated serum: No growth. Broth : Superficial pellicle composed of pink colonies with white aerial mycelium ; moderate flocculent sediment . Glucose broth : Small sediment of fine flocculi ; later pellicle composed of small pink colonies; superficial skin entire and salmon-colored in 16 days. Synthetic glycerol solution : Round pink disc-like colonies on surface and tenuous white wispy growth in suspension and sediment; after 20 days, surface colonies bearing white aerial mycelium extending 2 cm up tube. Synthetic sucrose solution : Minute white colonies in suspension and sedi- ment in 3 days; thin dust-like pellicle in 10 days; some surface colonies with white aerial mycelium in 17 days. Milk : Red surface skin ; solid coagulum. Litmus milk: Red surface growth, no change in liquid; after 4 weeks, liquid decolorized, opaque. Potato plug: Abundant growth, small colonies, mostly confluent, entirely cov- ered with pale pink aerial mycelium ; growth becomes membranous, consider- ably buckled ; later superficial colonies with pink aerial mycelium on liquid at base of tube, bottom growth of round white colonies. Starch not hydrolyzed. Source: From lesions in goats. 8. Nocardia pretoriana Pijper and Pullinger. (Pijper and Pullinger, Jour. Trop. Med. Hyg., 30, 1927, 153; Actino- myces preiorianus Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 38.) Named for Pretoria in South Africa. Description from Erikson. (Med. Res. Council Spec. Rept. Ser. 203, 1935, 30). Minute fiat colonies are formed con- sisting of angularly branched filaments, and bearing a few short straight aerial hyphae; later the growth becomes spreading and extensive, the slipping of the branches is well marked and the aerial hyphae are divided into cylindrical conidia. Slightly acid-fast. Gelatin: A few colorless flakes. No liquefaction. Glucose agar : Pale buff umbilicated and piled up colonies. Gl3^cerol agar: Piled up pink mass, very scant white aerial mycelium at margin. FAMILY ACTINOMYCETACEAE 901 Ca-agar : Yellowish wrinkled coherent growth with white aerial mycelium on apices and at margin. Coon's agar: Colorless mostly sub- merged growth, scant white aerial myce- lium. Dorset's egg medium: A few round colorless colonies in 3 days ; after 3 weeks, irregular raised pink mass, warted ap- pearance, moderate degree of liquefac- tion. Serum agar: Raised, convoluted, slightly pinkish growth. Inspissated serum: No growth. Broth: Moderate quantity of Hakes and dust-like surface growth. Synthetic sucrose solution: A few colorless flakes on the surface, lesser bottom growth. Milk: Yellowish surface growth; solid coagulum in one month; later, partly digested, pale pink growth up the wall of the tube. Litmus milk : Colorless surface growth, liquid blue ; becoming hydrolyzed and decolorized. Potato plug: Small raised pale pink colonies with white aerial mj'celium; after 2 months, plug and liquid dis- colored, growth dull buff, dry and con- voluted at base, round and zonate at top of slant, white aerial mycelium, surface and bottom growth on liquid. Source : From a case of mycetoma of the chest wall in a South African native. Habitat : Human infectioas so far as known. 9. Nocardia pulmonalis (Burnett) comb. nov. (Actinomyces pulmonalis Burnett, Ann. Rept. N. Y. State Vet. Coll., 1909-1910, 167.) From Latin pulmo, lung. Gram-positive mj^celium breaking up readily into oval-shaped conidia. Acid- fast, especially in early stages of growth. Gelatin: Small, whitish, spherical colonies ; edges of colony becoming chalky white; limited liquefaction. Agar: Moist, raised growth in the form of small, spherical colonies. Glucose agar: Dull, whitish, convo- luted growth. Broth: Delicate, translucent film on surface, becoming corrugated with some whitish, spherical colonies in medium. Milk: Colonies on the surface of the medium; milk is coagulated in a few days, later digested. Potato : Lu.xuriant growth in the form of small, translucent, round colonies; becoming colored lemon-yellow; later, growth becomes convoluted or folded with chalky white aerial mycelium, color of plug brownish. Non-pathogenic for rabbits and guinea pigs. Aerobic . Source : From the lungs of a cow. Habitat: Bovine infections so far as known . 10. Nocardia parafBnae (Jensen) comb, nov. {Proaclinomyces paraffinae Jensen, Proc. Linn. Soc. New So. Wales, 56, 1931, 362.) From INI. L. paraffina, paraffine. In agar media, the organism initially forms an extensive mycelium of long, richly-branching hyphae, 0.4 to 0.5 mi- cron thick. After 5 to 6 days, at room temperature, numerous end branches swell to about double thickness, become more refractive, e.xhibit fine incisions along their external contours, and divide into oval, spore-like elements, 0.8 to 1.0 by 1.2 to 1.5 microns. This process of di- vision starts at the tips of the swollen branches and proceeds basipetally until most of the hyphae appear divided. Pri- mary septa have not been seen in the hyphae. A similar process of division takes place in liquid media, where also the filaments often fall into fragments of variable length. The spore-like elements, but not the undivided filaments, are markedly acid-fast. The aerial myce- lium consists of rather short, straight, not very much branched hyphae, 0.4 to 902 MANUAL OF DETERMINATIVE BACTERIOLOGY 0.6 micron thick, which never show any differentiation into spores. Gelatin: No liquefaction. Sucrose agar: Very scant growth. Thin colorless veil, sometimes with a trace of white aerial mycelium. Glucose agar: Fair growth. Vegeta- tive mycelium flat, growing into me- dium; pale ochre-yellow to orange, with raised outgrowths on the surface. Growth of a crumbly consistency. Scant, white, aerial mycelium. Nutrient agar: Slow but good growth. Vegetative mycelium superficial, some- what raised, ochre-yellow, hard, but with a loose, smeary surface. Aerial myce- lium scant, small white tufts. No pig- ment. Potato: Fair growth. Vegetative mycelium granulated, first pale-yellow, later deep ochre-yellow to orange. Scant, white, aerial mycelium. No pig- ment. Liquid media (milk, broth, synthetic solutions) : Small, round granules of various yellow to orange colors, firm, but can be crushed into a homogeneous smear. In old broth cultures a thick, hard, orange to brownish surface pellicle is formed. Sucrose not inverted. Starch not hydrolyzed. Cellulose is not decomposed. Nitrates are not reduced to nitrites. Milk is not coagulated or digested. Final reaction in glucose NH4CI solu- tion, pH 4.6 to 4.4. All strains show a marked power of uti- lizing paraffin wax as a source of energy. Source : Isolated from soil. Habitat : Soil. 11. Nocardia minima (Jensen) comb, nov. {Proactinomyces minimus Jensen, Proc. Linnean Soc. New So. Wales, 56, 1931, 365.) From Latin minimus, very small . Filaments and rods : 0.4 to 0.6 by 2 to 10 microns. In older cultures mostly short rods, frequently V, Y, swollen forms, or cocci. Irregularly stained with ordinary dyes, generally show bars and bands. Generally a few cells from cul- tures are acid-fast, most are not acid- fast. Gram-positive. Gelatin stab : Filiform, granulated, cream-colored growth. No liquefaction. Agar : Slow growth, raised, folded, with finely myeloid margins. At first color- less, after 6 to 8 weeks flesh pink or coral pink. Potato : Growth slow, after 6 to 8 weeks abundant, spreading, much raised, finely wrinkled, coral pink. Paraffin is utilized. Optimum temperature 22° to 25°C. Distinctive characters : Closely re- sembles Nocardia corallina but differs in the much slower growth and the smaller size of the cells. Source : From soil in Australia. Habitat: Soil. 12. Nocardia corallina (Bergey et al.) comb. nov. (Bacillus mycoides corallinus Hefferan, Cent. f. Bakt., II Abt., 11, 1904, 459; Serratia corallina Bergey et al., Manual, 1st ed., 1923, 9Z;Streptothrix corallinus Reader, Jour. Path, and Bact., 29, 1926, 1 ; Mycobacterium agreste Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 84; Actinomyces agrestis Bergey et al., Manual, 3rd ed., 1930, 472; Pro- actinomyces agrestis Jensen, Proc. Lin- nean Soc. New So. Wales, 56, 1931, 345; Proactinomyces corallinus Jensen, ibid., 57, 1932, 364.) From Latin corallinus, coral red. Description from Gray and Thornton {loc. cit.), Jensen {loc. cit.) and Bynoe (Thesis, McGill University, Montreal, 1931). Branching rods, generally curved, 1 to 1.5 by 3 to 10 microns. In older cultures generally shorter rods and cocci. Non- motile. Not acid-fast. Gram-positive. Gelatin colonies : Round, convex, smooth, pink, shining, edge filamen- tous. Deep colonies: Burrs. FAMILY ACTINOMYCETACEAE 903 Gelatin stab: Nailhead; line of stab arborescent. No liquefaction. Agar colonies : Round, convex, or um- bonate, smooth, pink, shining or matte; border lighter, edge filamentous or with arborescent projections. Deep colonies: Burrs, or lens-shaped, with arborescent projections. In their verj'' early stages colonies consist of branching filamentous rods. As the colony grows, the cells in the interior break up into short rods and cocci which eventually form the mass of the colony. Cells on the outside remain filamentous, giving the colony a burr- like appearance, and often forming long arborescent processes. Agar slant: Filiform, convex, smooth, pink, shining or matte; arborescent or with projections from undulate border. Litmus milk: Alkaline. Reddish pel- licle. Glycerol potato: Filiform, raised, dry, wrinkled, yellomsh-brown to coral red. Broth: Usuallj^ turbid. Pink scum. Dorset's egg medium: Filiform, raised, dry, wrinkled, orange. Loeffler's medium: Similar to growth on Dorset's egg medium, but pink. Nitrites produced from nitrates. Acid from glycerol and glucose with some strains. Xo acid or gas from su- crose, maltose or lactose. Phenol and m-cresol are utilized. Some strains utilize naphthalene. (Gray and Thornton.) Some strains utilize phenol or m-cresol (Jensen). Optimum pH 6.8 to 8.0. Optimum temperature 22" to 25°C. Distinctive characters : Soil organism forming M ycobacterium-like colonies after 2 to 4 days on simple media. Pale pink chromogenesis . Xailhead growth in gela- tin stab. Branching rods and short filaments. Source : Seventy-four strains isolated from soils in Great Britain and Australia. Habitat: Soil. 13. Nocardia globerula (Gray) co^nb. nov. (Mycobacterium globerulum Gray, Proc. Roy. Soc. London, B, lOS, 1928, 265; Proaciinomyces globerulus Reed, in Manual, 5th ed., 1939, 838.) From Latin globus, a sphere. Description from Gray (loc. cit.) and from Bynoe (Thesis, McGill University, Montreal, 1931). Curved rods and fUaments : 1 by 2 to 9 microns, with many coccoid cells, es- pecially in old cultures. Rods and fila- ments are frequently irregularly swollen. Xot acid-fast. Capsules may be present. Gram-positive. Gelatin : After 19 days surface colonies irregularly round, 1 to 2 nma in diameter, convex, light buff, smooth, shining; edge entire. Deep colonies : Round, with en- tire edge. Gelatin stab : After 8 days nailhead, irregularly round, convex, pinkish-white, smooth, shining; line of stab erose. Agar : After 4 days surface colonies ir- regularly round, 3 to 5 mm in diameter, convex, white, smooth, shining; edge un- dulate, erose. After 7 days, more con- vex and of a watery appearance. Deep colonies : After 4 days, lens-shaped. Agar slant : After 3 days, filiform, fiat, watery; edge irregular. Nutrient and peptone broth : Turbid with \'iscous suspension. Indole not formed. Litmus milk: Alkaline. Glycerol potato: After 24 hours, fili- form, moist, smooth, pale pink. Dorset's egg medium: After 2 weeks, spreading, raised, moist, orange -colored. Loeffler's medium: Growth jas on Dorset's egg medium, but salmon-colored. Xitrites not produced from nitrates. Xo acid from glucose, lactose, maltose, sucrose or glycerol. Phenol is utilized. Indole agar : Blue crystals of indigotin formed. Optimum temperature 25° to 28°C. Optimum pH 6.8 to 7.6. Distinctive characters : This organism resembles most closely Nocardia corallina. It is distinguished by pro- 904 MANUAL OF DETERMINATIVE BACTERIOLOGY ducing a more watery type of surface growth, more nearly entire deep colonies and more particularly by the production of indigotin from indol. Source: From soil in Great Britain. Habitat: Presumably soil. 14. Nocardia salmonicolor (den Dooren de Jong) comb. nor. {M ycobacterium salmonicolor den Dooren de Jong, Cent, f. Bakt., II Abt., 71, 1927, 216; Flavo- hacterium salmonicolor Bergey et al., Manual, 3rd ed., 1930, 157; Proactino- myces salmonicolor Jensen, Proc. Linnean Soc. New So. Wales, 57, 1932, 368.) From Latin salmo, salmon and color, color. Closely related to Nocardia corallina. On glucose -as paragine-agar after 18 to 24 hrs., long branching rods are formed, 1.0 to 1.3 microns in thickness, with small refractive granules of aerial my- celium, sometimes stretching into quite long filaments; after 2 to 3 days small definite mycelia are present, and after 5 to 6 days these have largely divided into short rods and cocci; the colonies have the same burr-like appearance as those of Nocardia corallina. Many cells at the edge of the colonies show, after 3 to 4 days, club- or pear-shaped swellings, up to 2.5 to 3.0 microns in width; after 5 to 6 days, many of these swollen cells are seen to germinate with the formation of two more slender sprouts. {0rskov, Investigations into the Morphology of the ray fungi. Copenhagen, 1923, 82, gives an almost identical picture of Streptothrix rubra; it is questionable, indeed, whether these two organisms are not really identical.) Gelatin: At 20'' to 22°C, scant arbores- cent growth in stab; small wrinkled orange surface colony. No liquefaction. Glucose-asparagine-agar : Good growth, restricted, rather flat, edges lobate, sur- face warty, glistening, first pale orange, later ochre-yellow; consistency crumbly. After 5 to 6 weeks the growth is paler with manj' small round raised j'ellow secondary colonies. Glucose-nutrient agar: Excellent growth, spreading, fiat, dense, edges lo- bate, surface folded, glistening, yellow, gradually changing to deep orange-red. Nutrient broth : Fair growth ; thin pel- licle and granular sediment, at first cream-colored, later red; broth clear at first, slightly turbid after 3 w'eeks. Milk : Good growth ; pellicle of small cream-colored granules after 2 days, later a thick orange sediment. Not coagu- lated, but appears slightly cleared after 5 weeks, the reaction becoming alkaline. Potato. Good growth, raised, warty, crumbly, glistening, at first buff, chang- ing to orange and finally to almost blood - red. Indole not formed. Nitrites produced from nitrates. Nitrate, ammonium salts, asparagine and peptone are utilized almost equally well with glucose as source of carbon, although the growth is most rapid with peptone. Sucrose not inverted, although readily utilized with sodium nitrate as a source of nitrogen. Paraffin readily utilized as a source of carbon . Phenol not utilized. No acid from glucose or glycerol. Starch not hydrolyzed. No growth in oxygen-free atmosphere. Source : Isolated from soil by means of an ethylamine enriched medium, at 37°C. Habitat: Probably soil. 15. Nocardia rubropertincta (Heffer- an) comb. nov. (Butterbacillus, Grass- berger, Mi'inch. med. Wochnschr., 46, 1899, 343; Bacillus rubropertinctus Hef- feran. Cent. f. Bakt., II Abt., 11, 1903, 460; Serratia rubropertinctus Bergey et al.. Manual, 1st ed., 1923, 96; Mycobac- terium rubropertinctum Ford, Textb. of Bact., 1927, 255; Proactinomyces ru- bropertinctus Reed, in Manual, 5th ed.. FAMILY ACTINOMYCETACEAE 905 1939, 835.) From Latin, colored very red. Biittner (Arch. Hyg., 97, 1926, 17) regards Mijcohacterium cos as probably identical with Mycobacterium ruhrum Sohngen (Cent. f. Bakt., II Abt., 37, 1913, 599), Grassberger's organism {loc. cit.), Hormann and Morgenrot's organ- ism (Hyg. Rundsch., 7, 1898, 229), and Weber's organism (Arb. a. d. k. Gesund- heitsamte, Berlin, 19, 1903). To this list Lehmann and Neumann (Bakt. Diag., 7 Aufl., 2, 1927, 764) also add the organism of Ascher (Ztschr. f. Hyg., 32, 1899, 329) and the butter bacillus of Aujeszky (Cent. f. Bakt., I Abt., Orig., 31, 1902, 132). Jensen (Proc. Linnean Soc. New So. Wales, ^9, 1934, 32) regards the following organisms as probably identical : Bac- terium, ruhrum Migula (Syst. d. Bakt., 2, 1900, 488) a preliminary description of which is given by Schneider (Arb. bakt. Inst. Karlsruhe, 1, Heft 2, 1894, 213); probably this is also the organism re- ferred to by Haag (Cent. f. Bakt., II Abt., 71, 1927, 35) as Bacterium ruhrum; and Mycobacterium ruhrum Sohngen (loc. cit.). Description taken from Grassberger (Zoc. c;7.), Hefferan (loc. cit.) and Jensen (loc. cit.). Small rods: 0.3 to 0.9 by 1.5 to 3.0 microns. Cells in 18 to 24 hour agar culture in beautiful angular arrangement, after 2 to 3 days nearly coccoid, 0.6 by 0.8 micron. Tendency for branching on glycerol agar after 2 to 3 days, but branch- ing does not occur commonly though granules of aerial mycelium are sometimes seen (Jensen). Not acid-fast (Grass- berger). Acid-fast (Hefferan). Vari- able (Jensen). Non-motile. Gram- positive. Gelatin colonies : Irregular with crenate margin and folded surface. Coral red. Gelatin stab : Surface growth like the colonies. Growth in stab at first thin, then granular to arborescent with chro- mogenesis. No liquefaction. Agar colonies: Small, granular, be- coming pink to red depending on com- position of agar. Agar slant : Dry, lustreless (R) to glistening (S), pink to vermillion red. Broth : Faint uniform turbidity with salmon-pink pellicle (in scales) which is renewed on surface as it settles to form a red sediment (Hefferan, Jensen). Litmus milk: Thick, fragile, dull coral red surface scales and sediment. Un- changed (Hefferan) to alkaline and somewhat viscid after 3 to 4 weeks (Jensen). Potato : Slow but excellent intensive red growth becoming dull orange (Jensen). Nitrites not produced from nitrates; nitrates, ammonia and asparagine are al- most as good sources of nitrogen as pep- tone (Jensen). Benzine, petroleum, paraffin oil and paraffin are utilized as sources of energy (Sohngen). No action on manganese dioxide (Sohngen, Cent. f. Bakt., II Abt., 40, 1914, 554). Optimum pH 6.8 to 7.2. Growth stops at pH 4.9. Temperature relations: Grows well be- tween 20"" and 37°C (Jensen). Aerobic to facultative anaerobe. Distinctive characters: Mi/cobacie- rzMm-like colonies with coral to vermillion red chromogenesis on asparagine agar, potato, gelatin and other media. Short rods, seldom forms filaments. Generally not acid-fast. Source : Six cultures isolated from but- ter (Grassberger). Several cultures iso- lated from soil in Holland (Sohngen) and Australia (Jensen). Two cultures as contaminants in tuberculin flasks (Hagan, Breed). Habitat : Probablj'' widely distributed in soil. 16. Nocardia rubra (Krassilnikov) comb. nov. (Proaclinomyces ruber Kras- silnikov, Bull. Acad. Sci. U. S. S. R., No. 1, 1938, 139.) From Latin ruber, red. 906 MANUAL OF DETERMINATIVE BACTERIOLOGY Mycelium produced at first but soon breaks up into rods and cocci . The latter multiply by fission, cross-wall formation and budding. Cells are Gram-positive, weakljr acid-fast. Gelatin : No liquefaction. Colonies smooth or folded and rough, shiny or dull, dark red color. Pigment belongs to the carotinoids, does not diffuse into the medium. Milk: No coagulation or peptonization. Sucrose not inverted. Starch not hydrolyzed. No growth on cellulose. Readily assimilates fats and paraffin, and, to a less extent, wax. Various strains of this organism may vary considerably from type. Habitat : Soil. 17. Nocardia coeliaca (Gray and Thornton) comb. nov. {Mycohactcriuni coeliacum Gray and Thornton, Cent. f. Bakt., II Abt., 73, 1928, 88; Flavobacie- rium coeliacum Bergey et al., Manual. 3rd ed., 1930, 156; Proactinomyces co- eliacus Reed, in Manual, 5th ed., 19-39, 836.) Description from Gray and Thornton {loc. cil.) and from Jensen (Proc. Lin- nean Soc. New So. Wales, 56, 1931, 201). Short, curved, uneven-sided rods : 0.8 by 5 microns with occasional filaments up to 10 to 12 microns long, frequently beaded, occasionally swollen or branched. Coccoid forms 0.8 to 1.2 microns in di- ameter are common, especially in older cultures. Stain readily. Not acid -fast or occasionally slightly acid-fast . Gram- positive. Gelatin colonies : After 12 days, irregu- lar, raised, white, rugose, dull, edge en- tire. Deep colonies : Irregular, smooth or slightly broken. Gelatin stab: Convoluted, buff-white to yellowish, dull; below surface the growth forms many irregular hollow lobes, giving a glistening appearance, to a depth of 3 to 4 mm. Agar colonies: After 11 days, less than 1 mm in diameter, round or irregular, raised, white, resinous, edge irregular, burred. Deep colonies : Irregularly round or oval, edge slightly broken. Agar slant : Filiform, convex, white, rugose, resinous, edge undulate. Broth: Turbid. Litmus milk: Slightly alkaline after 5 to 7 days. Glycerol potato: After 2 days, dry, crumpled, orange, becoming brown after about 10 da5^s. Dorset's egg medium: Raised, smooth, moist, verrucose, buff-colored. Loeffler's medium: After 10 days, slight growth, dry, granular, pale buff- colored. Nitrites are not produced from nitrates. No acid from glucose, lactose, sucrose or glj'cerol. Phenol is utilized. Optimum pPI 7.6 to 8.0. Optimum temperature 22° to 25°C. Distinctive characters : Differs from the previously described members of the genus in the absence of chromogenesis. Forms hollow lobes in deep gelatin cul- tures. Cells are rods, seldom filaments. Source : From soil in Great Britain and Australia. Habitat : Presumably soil. IS. Nocardia transvalensis Pijper and Pullinger. (Pijper and Pullinger, Jour. Trop. Med. Hyg., 30, 1927, 153; Ac- tinomyces transvalensis Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 46.) Named for the Transvaal, a state of South Africa. Description from Erikson (Med. Res. Council Spec. Rept. Ser. 203, 1935, 28). Initial mycelium unicellular, but with the central branch frequently broader and showing dense granular refractile contents, small colonies quickly covered with aerial mycelium, the straight aerial hyphae in some cases becoming clustered into irregular spikes, colorless drops are exuded and a pink coloration produced in FAMILY ACTINOMYCETACEAE 907 the densest part of the growth on s\ni- thetic glj^cerol agar. Angular branching with division of the substratum filaments can be seen, the aerial hyphae also being irregularly segmented. Acid-fast. Gelatin : Poor gi'owth, a few irregular colorless flakes. Xo liquefaction. Agar: No growth. Glucose agar: Raised, granular, pink colonies with white aerial mycelium. Glycerol agar : Small pink coiled masses with thin white aerial mycelium. Potato agar: No growth. Coon's agar: Colorless growth with liberal white aerial mycelium. Dorset's egg medium : Small irregularly raised and coiled dull pink mass. Serum agar: Very poor growth. Inspissated serum : Scant colorless flaky growth; later a minute tuft of pale pink aerial mycelium. Broth: Moderate flaky growth. Synthetic sucrose solution : Poor growth, a few flakes on surface, a few at bottom. Potato plug: Dry, raised, convoluted, pink growth with white aerial mycelium in one month; dull, pink, brittle surface colonies, with paler aerial mycelium floating coherently on liquid at base in 2 months . Milk: No change. Starch not hydrolyzed. Source : From a case of mycetoma of the foot in South Africa. Habitat : Human infections so far as known. 19. Nocardia mesenterica (Orla-Jen- sen) comb. nov. (Microbacterium mesen- tericum Orla-Jensen, The Lactic Acid Bacteria, 1919, 181 j Proactinomyces mes- entericus Jensen, Proc. Linnean Soc. New So. Wales, 57, 1932, 373.) From Greek mesenterium, mesenter3^ Extensive mj-celium composed of richly branching hyphae of a somewhat variable thickness, 0.4 to O.S micron; no aerial hyphae are seen. With increasing age the hyphae divide into fragments of varying size and shape, partly diphthe- roid rods, but no real cocci. There is, particularly in richer media, a tendency to form large, swollen, fusiform to almost spherical cells, up to 3.5 microns in diameter. These may stain intensely with carbol fuchsin ; when transferred to fresh media, the}' germinate and produce a new mycelium. Gelatin: Good growth; finely arbores- cent, cream-colored growth in the stab; raised, folded, pale-yellow, surface colony. No liquefaction. Glucose-asparagine-agar : Fair growth, narrow, raised, granular, very pale yel- low, glistening; condensation water clear, with small granules. At 30°C only scant growth consisting of small irregular white granules, growing deeply down into the agar. Glucose-nutrient-agar : Good growth, restricted, with undulate edges, surface with high transverse folds, cream- colored; the consistency is firm and car- tilaginous after 2 daj's, later looser and more brittle. Growth at 28° to 30°C rather scant; smooth, soft, glistening, cream-colored smear, Sabouraud's agar: E.xcellent growth, spreading, at first flat and smooth, pale straw-yellow, perfectly hard and car- tilaginous, later raised and strongly folded, of a loose, curd-like consistency, bright lemon-yellow. Growth at 28° to SO^C onlj' fair, restricted, folded, cream- colored, soon becoming soft and smeary. Potato: Scant growth; restricted, soft, cream-colored smear. Broth: Good growth; voluminous, flaky, whitish sediment ; broth clear. Milk: At 28° to 30°C, small cream- colored granules along the tube ; the milk undergoes no visible changes within 4 weeks. No proteolytic action. Indole not formed. Sucrose is inverted. Starch is hydrolyzed. Cellulose is not decomposed. Nitrates are reduced to nitrites. No growth in o.xygen-free atmosphere. 908 MAMAL OF DETEKMlAATiVK BACTERIOLOGY Nitrogen is utilized as sodium nitrate, ammonium phosphate and asparagine, although tliese are inferior to peptone as sources of nitrogen. Source: Fermented beets. 20. Nocardia flava (Krassilnikov) comb. tiov. (Proactinoniyces flavus Kras- silnikov, Bull. Acad. Sci. U. S. S. R., No. 1, 1938, 139.) From Latin flavus, yellow. Cells at first filamentous, 0.7 to 0.8 micron in diameter; after 2 to 3 days broken into long rods and then into cocci 0.7 micron in diameter. No spores, some strains form chlamydospores. Cell multiplication by fission, cross-wall for- mation, rarely by budding. Cells Gram-positive; not acid-fast. Gelatin: No liquefaction. Synthetic agar colonies : Bright yellow or gold color. Meat peptone media : Dirty yellow pigmentation. Agar colonies : Pigment bright yellow or gold color on synthetic media, dirty yellow on meat peptone media. Pigment not soluble in medium. Surface of colony somewhat shiny or rough and folded, of a dough-like consistency. Milk: No peptonization or coagulation. Sucrose weakly inverted. Starch is hydrolyzed. Does not grow on paraffin and wax but makes weak growth on fat. Habitat : Soil, not common. 21. Nocardia viridis (Krassilnikov) comh. 710V. {Proaclinomyces viridis Krassilnikov, Bull. Acad. Sci. U. S. S. R., No. 1, 1938, 139). From Latin viridis, green. Mycelial cells often branching, 0.7 to 0.8 micron in diameter with cross-wall; after 5 to 7 days the cells break up into rods 5 to 15 microns long. Cocci not observed. Cells multiply by fission, seldom by budding. Spores not formed. Cells Gram-positive, not acid-fast. Gelatin: No liquefaction. Colonies colored dark green. Pigment not soluble in medium, in water or in organic solvents. Surface of colony somewhat shiny. On potato, rough, much folded, broken up into small col- onies. Milk: No peptonization or coagulation. Sucrose readily inverted. Starch weakly hydrolj^zed. Grows well on fats and paraffin and less on wax. Habitat: Soil. 22. Nocardia citrea (Krassilnikov) comb. nov. {Proaclinomyces citrous Krassilnikov, Bull. Acad. Sci. U. S. S. R., No. 1, 1938, 139.) From M. L. citreus, lemon yellow. Mycelium in young cultures consists of ver3' fine threads 0.3 to 0.5 micron in diameter. After several days the cells break up into short rods 0.5 by 1.5 to 5 microns and into cocci 0.3 to 0.5 micron in diameter. Multiplies by fission and bud formation ; spores not formed. Cells not acid -fast. Gelatin: Liquefaction. Colonies: Yellow -green, usuallj^ rough and folded. Milk: Coagulation and peptonization. Sucrose is inverted. Starch is hydrolyzed. Weak growth on fat. No growth on paraffin or wax. Habitat : Soil and water. 23. Nocardia madurae (Vincent) Blanchard. {Streptothrix madurae Vin- cent, Ann. Inst. Past., 8, 1894, 129; Blanchard, Bouchard's Traite de Path. Gen., 2, 1896, 868; Oospora madurae Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 388; Actinomyces madurae Lachner-Sandoval, Ueber Strahlenpilze, 1898, 64; Cladothrix madurae Mace, Trait<5 Pratique de Bact., 4th ed., 1901, 1090; Discomyces madurae Gedoelst, Champ. Pai'as. Homme et Animaux, Paris. 1902. 169. ) Named for the disease FAMILY ACTINOMYCETACEAE 909 Madura foot, with which this organism is associated. Oospora indica Kant hack (Ivanthack, Jour. Path, and Bact., /, 1893, 140; No- cardia indica Chalmers and Christo- pherson, Ann. Trop. Med. and Parasit., 10, 1916, 231 ; Discomyces indicus Neveu- Lemaire, Precis de Parasitol. Hum., 5th ed., 1921, 42; Actinomyces indicus Brumpt, Precis de Parasitol., Paris, 4th ed., 1927, 1196) is regarded by some authors as identical with Nocardia madurae Blanchard. If this is estab- lished, then the correct name of the organism is Nocardia indica (Kanthack) Chalmers and Christopherson. The species described under the name Actinomyces madurae in previous editions of Bergey's Manual is definitely not the true causative agent of the disease and is probably a contaminant carried as a cul- ture of this species. Morphology in tissues, growth in form of granules consisting of radiating actinomycosis. In cultures, initial branched mycelium fragmenting into rod -shaped and coccoid bodies. No aerial mycelium or spores. Not acid- fast. Gelatin: Growth scant, whitish; no liquefaction. Gelatin colonies : Round, glistening, at first white, then buff to rose or crimson. Pigment production is irregular and un- predictable. Occasionally red soluble pigment is produced. Growth eventu- ally wrinkled. No aerial mycelium. Potato: Wrinkled friable growth; buff- colored, sometimes red. Broth : Growth as a floccular sediment. Milk: No change, or slight slow pep- tonization. Diastatic (?) action. Not pathogenic for the usual laboratory animals; pathogenic for monkeys (Mus- grave and Clegg, Philippine Jour. Sci., Ser. B., Med. Sci., 3, 1908, 470). Habitat : Cause of some cases of Madura foot. 24. Nocardia lutea Christopherson and Archibald. (Christopherson and Archi- bald, Lancet, 2, 1918, 847; Actinomyces luteus Brumpt, Precis de Parasitol., Paris, 4th ed., 1927, 1206.) From Latin luteus, yellow. Description from Erikson (Med. Res. Council Spec. Rept. Ser. 203, 1935, 30). Initial elements swollen and seg- mented, giving rise to irregular spreading polymorphous colonies composed of cells of all shapes and sizes with markedly granular contents. Later more mono- morphous, the filaments being arranged in angular apposition. Sometimes (e.g., on synthetic glycerol agar) the segments are so granular as to appear banded. On potato agar, small filamentous colonies are formed with irregular angular branch- ing and bear a few isolated short straight aerial hyphae. Gelatin : Pale pink wrinkled growth on wall of tube and colorless punctiform and stellate colonies in medium ; no liquefac- tion. Agar: Abundant, coherent, moist, pink membranous growth with round discrete colonies at margin ; after 3 weeks colorless fringed margin, round confluent portion. Glucose agar : Scant reddish smeary growth. Glycerol agar : Yellowish-pink, wrinkled membrane. Potato agar : Coherent pink moist growth, centrally embedded with small round discrete colonies at margin. Dorset's egg medium: Poor growth, dull pink, spreading. Serum agar : Confluent granular pink membrane. Broth : Pink flakes and surface growth. Inspissated serum : Raised convoluted pink mass; becoming orange, much wrinkled, scalloped margin. Synthetic sucrose solution : Red granules and abundant minute colorless colonies at bottom ; in 2 weeks a colorless dust-like surface pellicle. Glucose broth : Abundant, pinkish 910 ]VL\NUAL OF DETERMINATIVE BACTERIOLOGY flaky surface growth, breaking up easily and sinking to bottom. Litmus milk: Orange-red surface and bottom growth, liquid blue. Potato plug: Carrot-red, moist, thick, granular growth in bands, partly raised and with discrete round colonies ; sparse colorless very thin aerial mycelium at top of slant in 2 months. Source : From actinomycosis of the lachrymal gland. F 25. Nocardia blackwellii (Erikson) comb. nov. {Actinomyces blackwellii Erik- son, Med. Res. Counc. Spec. Rept. Ser. 203, 1935, 37.) Description from Erikson {loc. cit., p. 32). Initial elements short, rod-like, grow- ing out into longer forms sparsely branch- ing; small radiating colonies are produced with short straight aerial mycelium, frequently large round or ovoid cells are interposed in the irregularly segmented chains of cells, being sometimes isolated in company with 2 or 3 short filaments and sometimes terminal. Gelatin : Few colorless minute colonies along line of inoculation; after 30 days abundant colorless colonies to 10 mm be- low surface, larger pink-yellow surface colonies with white aerial mycelium; no liquefaction. Agar : Confluent wrinkled growth with small, round, pinkish, discrete colonies at margin. Glucose agar: Abundant, pale pink growth, small conical colonies, piled up, convoluted. Glycerol agar : Extensive, granular, ir- regular, thin, pinkish growth; after 40 days, a few discrete colonies with de- pressed margins, center piled up, pink. Serum agar: Smooth, cream, umbili- cated colonies, with submerged growth extending into medium in scallops 5 to 8 mm deep; a pale pink mass in 2 weeks, t Potato agar: Small, round, colorless colonies covered with white aerial my- celium; after 2 weeks colonies dull pink, submerged margins, few aerial spikes, moderate aerial mycelium at top of slant. Broth: Flakes, later innumerable minute colonies, some adhering to wall just above liquid level. Synthetic sucrose solution : Delicate, round, white colonies ; later abundant minute colonies in suspension, thick cream pellicle on surface and pink grains in sediment. Milk : Heav}' convoluted bright j^ellow surface pellicle, no coagulation. Litmus milk: Yellow surface growth, nrdlky sediment, liquid unchanged. Carrot plug: Small, round, smooth, cream-colored elevated colonies in 10 days; sparse stiff colorless aerial spikes in 16 da3\s ; abundantly piled up, con- voluted, ochreous growth in 25 days. Source : From hock joint of foal. 26. Nocardia cuniculi Snijders. (Snijders, Geneesk. Tijdsch. Med. Ind., 64, 1924, 47 and 75; Aclinoviyces cuniculi Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 32; not Actinomyces cuniculi Gasperini, Cent. f. Bakt., 15, 1894, 684; Actinomj/ces sumatrae Erikson, loc. cit., 32.) From Latin cuniculus, rabbit. Description taken from Erikson {loc. cit., p. 31). Large swollen cells give rise to ramify- ing filaments or to small chains of short thick segments which branch out into more regular hyphae; sometimes the ir- regular elements are beset with spiny processes before giving rise to typical long branching filaments ; later the pic- ture becomes more monomorphous and short straight aerial hyphae are borne, which presently exhibit irregular seg- mentation. Gelatin: Few flakes. No liquefaction. Agar: Small, round, elevated, cream- colored colonies, umbilical ed and radially wrinkled. Glucose agar: Minute, colorless colon- FAMILY ACTINOMYCETACEAE 911 ies ; becoming dull pink, partly confluent and piled up, fewstiif pink aerial spikes. Glycerol agar: Small round elevated cream-colored colonies, margins de- pressed; becoming smooth, discrete, yellowish. Dorset's egg medium: Scant pinkish smeary growi;h. Serum agar : Small, raised, cream- colored colonies, becoming confluent and piled up. Inspissated serum: Thick, colorless, ribbed membrane; no liquefaction. Broth : Small and larger cream-colored, scale-like surface colonies, abundant, flocculent bottom growth. Sj'nthetic sucrose solution; Thin sur- face pellicle, small colorless flakes, minute particles at bottom, scant growth. Milk : Heavj^ yellow growth attached to walls; solid coagulum in 1 month. Litmus milk : Yellow surface growth, liquid unchanged. Potato plug: Coral-pink, dry, granular growth, covered to a considerable extent with white aerial mycelium, piled up in center, discrete colonies at margin, pink surface pellicle on liquid and colorless colonies at base. Source: Infected rabbits. 27. Nocardia rangoonensis (Erikson) comb. nov. {Actinomyces rangoon Erik- son, Med. Res. Council Spec. Rept. Ser. 203, 1935, 37.) Description from Erikson (lac. cit., p. 33). Swollen round initial cells, giving rise to branching hyphae which segment and present slipping and angular arrange- ment; few short straight aerial hyphae, which later develop into a profusely branching long waving aerial mycelium. Non-acid-fast. Gelatin: Abundant minute colonies in depths and larger cream-colored ones on surface with white aerial mycelium; brown pigment surrounding growth. No liquefaction. Agar colonies : Round, lobate, umbili- cated, raised up, cream-colored to pale pink; later, medium discolored dark brown, colonies colorless. Glucose agar : Convoluted, coherent, cream-colored growth, medium dis- colored. After 23 days, wrinkled, bis- cuit-colored growth, colorless margin, border white aerial mycelium, medium dark brown. Glycerol agar: Dull, mealy, pink, wrinkled growth, scant white aerial myce- lium at top, medium slightly discolored. Coon's agar : Minute colorless colonies in streaks. Potato agar: Small, round, lemon- colored colonies, parth'' confluent, with white aerial mycelium; later medium dis- colored light brown; submerged grov\i;h greenish. Dorset's egg medium: Extensive color- less growth, pale pink aerial mycelium in center; later covered with a powdery pinkish-white aerial mycelium. Serum agar colonies : Irregular, small, elevated, cream-colored, frequently um- bilicated. Inspissated serum: Poor growth, small piled up pink mass . Broth: Abundant colorless growth, flocculent mass at bottom and pellicle at surface, medium slightly discolored. Synthetic sucrose solution : Small white colonies with pinkish tinge on surface, lesser bottom growth. Milk : Coagulation, j^ellow surface ring, becoming partly peptonized, liquid dis- colored dark brown, brownish growth up side of tube. Litmus milk: Colorless growth, liquid partly decolorized; coagulation; later partly digested. Carrot plug: Small round colorless colonies, velvety white aerial mycelium; in 2 months, piled up pink granular mass with warted prominences, marginal zone white aerial mycelium and thin all-over central aerial mycelium. Source : Human pulmonary case of streptothricosis. 912 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : Human infections so far as known. 28. Nocardia caviae Snijders. (Snidjers, Geneesk. Tijdschr. Ned. Ind., 64, 1924, 47 and 75; Actinomyces caviae Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 37.) From the generic name of the guinea pig. Description from Erikson {loc. cit., p. 32). Initial segmentation, producing ele- ments of appro.ximately even thickness arranged in angular apposition, and later long profusely ramifying threads with strongly refractile protoplasm. Aerial mycelium straight and branching, the aerial hyphae with occasional coiled tips divided into cylindrical conidia. Gelatin: A few colorless flakes. No liquefaction. Glucose agar: Piled up, convoluted, cream-colored to pale pink growth, white aerial mycelium. Glycerol agar: Scanty growth. Coon's agar: Colorless scant growth, partly submerged, white aerial mycelium. Potato agar : Colorless spreading growth with dense white aerial mycelium. Dorset's egg medium: Heavily corru- gated pale pink growth with submerged margin, dense white aerial mycelium in center ; after 3 weeks, colorless transpired drops. Serum agar: Pale pink wrinkled growth, partly submerged; after 4 weeks, piled up with scant white aerial myce- lium, medium discolored reddish-brown. Inspissated serum : Pale pink raised growth, coiled, white aerial mycelium. Broth : Cream-colored wrinkled surface pellicle extending up wall and breaking easily, moderate bottom growth, flaky, medium discolored. Synthetic sucrose solution: Round white colonies in suspension and attached to one side of tube, pink surface colonies with white aerial mycelium. Milk: Colorless surface growth, white aerial mycelium; coagulation. Litmus milk: Liquid blue, surface growth; after 1 month, white aerial my- celium, colorless sediment, liquid still blue. Potato plug: Small colorless colonies, white powdery aerial mycelium; later abundant raised pale pink confluent growth, discolored plug; after 2 months, raised buckled pink colonies with white aerial mycelium floating on liquid at base. Source : Infected guinea pigs, Sumatra. 29. Nocardia actinomorpha (Gray and Thornton) comb. nov. {Mycobacterium actinomorphum Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 88; Actino- myces actinomorphus Bergey et al., Manual, 3rd ed., 1930, 471; Proactino- myces actinomorphus Jensen, Proc. Lin- nean Soc. New So. Wales, 56, 1931, 363.) From Greek actis, ray and Latin morpha, shape, form. Description from Gray and Thornton (loc. cit.), Jensen {loc. cit.), and Bynoe (Thesis, McGill University, Montreal, 1931). Long branching filaments and rods : 0.5 to 0.8 by up to 10 microns. In older cul- tures rods 2 to 3 microns long generally predominate. On some media exten- tively branching hyphae occur. Readily stained. Not acid-fast. Gram-positive. Gelatin colonies : After 12 days, round, saucer-like, white, raised rim, edges burred. Liquefaction. Deep colonies: Burrs. Gelatin stab : After 8 to 14 days, sac- cate liquefaction, 5 to 8 mm. Agar colonies: After 11 days, round, 1 mm in diameter, convex, white, granular or resinous ; long arborescent processes from the edge. Deep colonies: Arbores- cent burrs ; processes about equal to diameter of colony. Agar slant : Filiform, raised to convex, white, rugose, dull; edge undulate, with strong tufted projections below surface. Broth: Turbid, or clear with white FAMILY ACTINOMYCETACEAE 913 Dorset's egg medium: After 2 weeks, raised, dry, smooth, salmon-buff growth. Loeffler's medium: After 2 days, smooth, moist, warty, salmon-colored growth. Litmus milk : Alkaline after 5 to 7 days. Glycerol potato: After 2 days, dry, wrinkled, pink to orange growth. Nitrites are produced from nitrates. No acid from glucose, lactose, sucrose or glycerol. Phenol and naphthalene are utilized. Optimum temperature 25° to 30°C. Optimum pH 7.8 to 8.5. Distinctive characters : Differs from Nocardia coeliaca in saccate liquefaction of gelatin. Long rods and filaments. Source : A few strains have been iso- lated from soil in Great Britain and Australia. Habitat : Presumably soil. 30. Nocardia flavescens (Jensen) comb. nov. (Proactinomyces flavescens Jensen, Proc. Linnean Soc. New So. Wales, 56, 1931, 361.) From Latin flavescens, becoming golden yellow. On media where a firm growth is pro- duced, the vegetative mycelium appears as long, branched, non-septate hyphae, 0.4 to 0.6 micron thick. In other media, as on nutrient agar and potato, septa are formed and the mycelium appears in preparations as fragments of very vari- able size, partly resembling highly branched mycobacteria. In several cases — for instance, on nutrient agar at 28° to 30°C, in 5 to 6 weeks old cultures in glucose broth, and in glucose NH4 CI solution — short elements assume swollen, fusiform to lemon-shaped forms. The aerial mycelium consists of fairlj' long hyphae of the same thickness as the vegetative hyphae, not very much branched, without spirals, often clinging together in wisps. A differentiation into spores is never visible by direct microscopic examination. Neither is this the case in stained preparations ; here the aerial hyphae break up into fragments of quite variable length, from 1.2 to 1.5 up to 10 to 12 microns, showing an irregular, granulated staining. Gelatin: Slow liquefaction. Sucrose agar: Good growth. Vegeta- tive mycelium superficially spreading, much raised and wrinkled, cracking, white to cream-colored, of a dry, but loose and crumbly, consistency. Aerial my- celium scant, thin, white. Faint yellow soluble pigment after 2 to 3 weeks. Glucose agar: Good growth. Vegeta- tive mj'celium superficial, wrinkled, honey-yellow, of a hard and cartilagin- ous consistency. Aerial mycelium thin, smooth, white. Yellow soluble pigment. Nutrient agar: Good growth. Vegeta- tive mycelium raised and much wrinkled, first dirty cream-colored, later dark yellowish-gray, of a soft, moist, curd-like consistency. No aerial mycelium. No pigment . Potato; Good to excellent growth. \'egetative mycelium much raised and wrinkled, first cream-colored, later yel- lowish-brown, soft and smeary. No aerial mycelium, no pigment. Glucose broth: Rather scant growth. Granulated, 3'ellowish sediment; no sur- face growth. Broth clear. No pigment. No acidity. Sucrose is inverted. Starch is hydrolyzed. Cellulose is not decomposed. Nitrates are reduced slightly or not at all with various sources of energy. Milk: Coagulated and slowly redis- solved with acid reaction. Final reaction in glucose -NH4CI solu- tion, pH 3.9 to 3.6. No growth under anaerobic conditions. Habitat: Soil. 31. Nocardia maculata (Millard and Burr) comh. nov. {Actinomyces macula- tus Millard and Burr, Ann. Appl. Biol., 13, 1926, 580; Proactinomyces maculatus Umbreit, Jour. Bact., 38, 1939, 84.) From Latin maculatus, spotted. Description taken from Umbreit. 914 MANUAL OF DETERMIXATIVE BACTERIOLOGY Filamentous organisms possessing a tough shiny colony which is cartilagin- ous, rarely producing an aerial mycelium, though in certain strains, it may occur frequently. Retains the mycelium form for long periods. Not acid-fast. Gelatin: Liquefaction. In the young colony an orange -yellow to orange-red intercellular pigment is produced on all media, which may or may not change to black as the culture ages. Milk: No digestion. Starch is hydrolyzed. Does not utilize paraffin. Habitat : Soil. 32. Nocardia rhodnii (Erikson) comb, nov. {Actinomyces rhodnii Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 37.) Named for the insect genus, Rhodniiis. Description from Erikson {luc. cil., p. 29). In early stages, the minute colonies are composed of hyphal segments arranged in angular apposition; the aerial mycelium being short and straight. Later the growth becomes extensive and spreading, made up partly of long, genuinely branch- ing filaments and partly of short segments exhibiting slipping branching, each giving rise to aerial hyphae. After 2 weeks the angular branching is very marked, deli- cate spreading herring-bone patterns being formed. Gelatin : Rapid liquefaction ; pale pink colonies in superficial pellicle and sedi- ment. Coon's agar : Colorless pinpoint colonies . Czapek's agar : Minute, colorless, round colonies. Glucose agar: Abundant, coral pink, convoluted, piled up growth. Glycerol agar: Extensive growth, dull pink colonies round and umbilicated, be- coming piled up and deeper coral ; later partly submerged. Dorset's egg medium: Salmon-pink, granular membrane; later piled up. Serum agar: Extensive, reddish, con- fluent mass, granular, tending to be piled up ; the medium around the growth shows reddish coloration in 2 weeks. Inspissated serum: Smooth, round, pale pink colonies, centrally depressed and irregularly coiled larger mass ; no liquefaction. Broth : Salmon-pink flakes in sediment and colonies on surface; after 2 weeks abundant growth, discoloration of me- dium. Glucose broth : Thin, pink, superficial pellicle, easil}' breaking up, and small flakes in sediment ; after 2 weeks abun- dant growth extending up tube. Synthetic sucrose solution: Colorless to pink colonies in superficial pellicle, and minute round white colonies co- herent in loosely branching mass in sediment. ]Milk : Bright orange growth ; medium unchanged. Potato agar: Abundant, pink growth, piled up, scant stiff white aerial myce- lium at top of slant. Source : From rcduvid bug, Rhodniiis prolixus. 33. Nocardia gardneri (Waksman) comb. nov. (Actinomycete, Gardner and Chain, Brit. Jour. Exp. Path., 23, 1942, 123; Proactinomijccs gardneri Waksman, in Waksman, Horning, Welsch and Wood- ruff, Soil Sci., 5J^, 1942, 289.) Named for Prof. Gardner who first isolated this organism. Gram-positive, branching mycelium. Gelatin : Cream-colored surface ring. Rapid liquefaction. Green to greenish- brown soluble pigment gradually diffuses through the liquefied portion. Nutrient agar: Cream-colored, ele- vated, lichenoid growth, soft, not leathery; no aerial mycelium; very faint brownish pigment. Glucose agar: Brownish, lichenoid FAMILY ACTIXOMYCETACEAE 915 growth, with wide, cream-colored edge; white to grayish aerial mycelium gradu- ally covering surface . Reverse of growt h yellowish; no soluble pigment. Glucose-asparagine agar: Aerial mj'ce- lium develops slowly. Tryptone broth: Growth occurs as small pellets at the base of the flask; later, a thin surface pellicle appears, which consists of a branching mycelium. Black pigment slowly produced. Litmus milk: Unchanged. Potato: Barnacle-like, brownish, spreading growth; no aerial mycelium. Medium brownish around growth. Indole not formed. Xo acid from glucose, lactose, maltose, mannitol, sucrose and dulcitol. Good growth at 25°C. Slow growth at 37°C. Distinctive character: Produces an antibiotic substance (proactinomycin) upon synthetic and organic media which is primarily active against various Gram- positive bacteria. Source : Isolated as an air contaminant at Oxford, England. * Appendix I : The following species probably belong to this genus. Many are incompletel}' described. Some of the species listed here ma}' belong in the genus Streptotnyces. Aciinoiiujces albiis ucidus Xeukirch. (Xeukirch, Inaug. Diss., Strassburg, 1902, 50; Aclinomyces albus var. acidus Xannizzi, in Pollacci, Tratt. ]\Iicopat. Umana, 4, 1934, 9.) From a case of keratitis. Actinomyces avadi Dodge. (Strepto- thrix madurae Koch and Stutzger, Ztschr. f. Hyg., 69, 1911, 17; not Streptothrix madurae Vincent, Ann. Inst. Past., 8, 1894, 129; Dodge, Medical Mycology, St. Louis, 1935, 729.) From a Madura foot in Egypt, case of Dr. Avad. Actinomyces bolognesii-chiurcoi (Vuil- lemin) Dodge. (Malbrachea bolognesii- chiurcoi Vuillemin, in Bolognesi and Chiurco, Archivi di Biol., 1, 1925; Dodge, :\Iedical Mycology, St. Louis, 1935, 766.) From ulcers on the thorax. Actinomyces cameli (Mason) Sartory and Bailly. (Streptothrix cameli Mason, Jour. Trop. Med. and Therap., 32, 1919, 34; Oospora cameli Sartory, Champ. Paras. Homme et Anim., 1923, 822; Sartory and Baillj', Mycoses Pulmo- naires, 1923, 253.) From pseudotubercu- losis lesions in a camel. Actinomyces canis (Rabe) Gasperini. (Discomyces pleuriticus Vachetta, Studi e ricordi clin. ]Milano, 1882; Pleu- romyces canis familiaris Rivolta, Giornali d. Anat. Fisiol. e Patol., 16, 1884, 4; Cladothrix canis Rabe, Berlin, tierarztl. Wochnschr., 1888, 65; Gasperini, Ann. 1st. d'lg. sper. Univ. Roma, 2, 1892, 222; Streptothrix canis and Actinomyces pleuriticus canis familiaris, quoted from Gasperini, Cent. f. Bakt., 15, 1894, 684; Leptothrix pleuriticus Piana and Galli-Vallerio, 1896, quoted from Xan- nizzi, in Pollacci, Tratt. Micopat. Lf^mana, 4, 1934, 37; Nocardia canis Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 255; Oospora canis Sartory, Champ. Paras. Homme et Anim., 1923, 821.) Rabe iso- lated this organism in two cases of phleg- mon and a case of peritonitis in dogs. Actinomyces citrocrevieus Xannizzi. (Mycobacteriiun diphtheriae avium Trin- cas, 1907, quoted from Xannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 50; Xannizzi, idem.) Froma disease in birds. Actinomyces dassonvillei Brocq- Rousseu. (Brocq-Rousseu, These Sci. * The appendix was originally pre])ared by Prof. S. A. Waksman and Prof. A. T. Henrici, May, 1943; it has been developed further by Mrs. Eleanore Heist Clise, Geneva, Xew York, August, 1945. 916 MANUAL OF DETERMINATIVE BACTERIOLOGY Nat. Paris, 1907; Nocardia dassonvillei Li^gard and Landrieu, Ann. d'Occulisti- que, 4^, 1911, 418; Disconujces dassonvillei Brumpt, Precis de Parasitologie, Paris, 2nd ed., 1913, 976.) Reported from a cervical abscess (Brumpt), from a case of conjunctivitis (Li^gard and Landrieu), and from grain (Pinoy, Bull. Inst. Past., 27,1913,923). Actinomyces dcrmatonomus Oxer. (Bull. Austral. Jour. Exp. Biol. Med. Sci., 6, 1929, 301, quoted from Dodge, iNIedical Mycology, St. Louis, 1935, 719; Oxer, 1930, quoted from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51.) From lesions on sheep in Australia. Actinomyces donnae Dodge. {Strepto- thrix sp. Donna, Ann. Ig. Sperim., 14, 1904, 449; Dodge, Medical Mycology, St. Louis, 1935, 745.) From sputum in a pulmonary infection. Actinomyces dori (Beurmann and Gou- gerot) Brumpt. {Sporotrichum, Dor, Presse M^d., 14, 1906, 234; Sporotrichum dori Beurmann and Gougerot, Ann. Derm. Syphiligr. IV, 7, 1906, 996; Dis- comyces dori Beurmann and Gougerot, Les Nouvelles Mycoses, 1909, 59; Rhino- cladium dori Neveu-Lemaire, Precis Parasitol., 1921, 84; Oospora dori Sartory, Champ. Paras. Homme et Anim., 1923, 770; Brumpt, Precis de Parasitol., 4th ed., 1927, 1206; Nocardia dori Vuillemin, EncyclopMie INIycologique, Paris, 2, 1931, 284.) Found in subcutaneous ab- scesses resembling sporotrichosis. Actinomyces hominis Berestneff. (Berestneff, Inaug. Diss., Moskow, 1897; not Actinomyces hominis Waksman, Soil Science, 8, 1919, 129; Nocardia hominis Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916; Discomyces hominis Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 984.) From a ca.se of pseudoactinomycosis. Actinomyces japonicus Caminiti. (Streptothrix sp. Aoyama and Miyamoto, Mitteil. Med. Fak. K. Jap. Univ. Tokio, 4, 1901, 231; abst. in Cent. f. Bakt., I Abt., Grig., 29, 1901, 262; Caminiti, Cent. f. Bakt., I Abt., Orig., 44, 1907, 198; Streptothrix japonica Pctruschky, in Kolle and Wasserman, Handb. d. path. Mikroorg., 2 Aufl., 5, 1913, 295; Discomyces japonicus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 984.) From a case of actinomycosis of the lungs. Actinomyces keratolytica Acton and McGuire. (Indian Med. Gaz., 65, 1930, 61 and 66, 1931, 65; Proactinomyces keratolyticus, author unknown.) Pro- duces cracked heels among the ryots of India. Actinomyces lepromatis de Souza- Araujo. (Compt. rend. Soc. Biol., 100, 1929, 937.) From a leproma, Brazil. Actinomyces levyi Dodge. (Acti- nomyces sp. Levj'^; Oospora sp. Sartory, Champ. Paras. Homme et Anim., 1923, 827; Dodge, Medical Mycology, St. Louis, 1935, 730.) From pus. Actinomyces micetomae Greco. {Streptothrix. micetomae argentinae P, Greco, in Durante, Segunda Observacion de Pie de Madura o Micetoma en la Re- publica Argentina, Thesis, Buenos Aires, 1911; Greco, Origine des Tumeurs, 1916, 759; Oospora micetomae Sartory, Champ. Paras. Homme et Anim., 1923, 783.) From a case of mycetoma pedis. Actinomyces miniiiius (LeCalve and r^Ialherbe) Dodge. (Oospora forme de Microsporum {Andouini var. eqtiinum), Bodin, Arch, de Parasitol., 2, 1899, 606; Trichophyton minimimi LeCalve and Malherbe, Arch, de Parasitol.; Micro- sporum minimum Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 993; Dodge, Medical Mycology, St. Louis, 1935, 728.) From ringworm of horse and dog. Actinomyces mucosus Basu. (Indian Med. Gaz., 78, 1943, 577.) From bron- chial actinomycosis. Actinomyces neddeni Namyslowski. (Streptothrix sp. zur Nedden, Klin. Monatsbl. f. Augenheilk., 45, 1907, 152; Namyslowski, Cent. f. Bakt., I FAMILY ACTINOMYCETACEAE 917 Abt., Orig., 62, 1912, 564.) I>om a case of keratitis. Actinomyces nodosus (Beveridge) Hagan. {Fusiformis nodosus Beveridge, Austral. Council Sci. and Indus. Res. Bui. 140, 1941, 56 pp.; Hagan, The In- fectious Diseases of Domestic Animals. Ithaca, Xew York, 1943, 312.) Con- sidered the primary cause of foot-rot of sheep. Also see Spirochaeta penortha. Actinomyces phenotolerans Werkman. (In Gammel, Arch. Derm. Sj^philol., 39, 1934, 286.) From granuloma in man. Actinomyces puntonii Lopez Ortega. (Lopez Ortega, Annali d'Igiene, Rome, ■H, 1934, 867; Asteroides puntonii Puntoni and Leonardi, Boll, e Atti d. R. Accad. Med. di Roma, 61, 1935, 94.) From a pulmonary abscess. Actinomyces purpureus Cavara. (Or- loff, Vestnik Ofth., 29, 1912, 653; Cavara, Micosi Occ, 1928, 99; not Actinomyces purpureus Killian and Feher, Ann. Inst. Past., 55, 1935, 620.) From a case of keratitis in Russia. Actinomyces ribeyro Dodge. (Hongo artrosporado Ribeyro, Ann. Fac. ]Med. Lima, 3, 1919, 1; Dodge, Medical Mycol- ogy, St. Louis, 1935, 735.) From a gener- alized infection on the arms, legs and chest of a patient in Peru. Actinomyces rodellae Dodge. {Strepto- thrix sp. Rodella, Cent. f. Bakt., I Abt., Orig., 84, 1920, 450; Dodge, Medical Mycology, St. Louis, 1935, 734.) From abscesses of the tooth and jaw. Actinomyces ruber (Kruse) Sanfelice. (Un Cladothrix cromogeno, Ruiz Casabo, Cronica medico-quirurgica de la Habana, 20, No. 13, 1894, 340; see Cent. f. Bakt., I Abt., 17, 1895, 466; Streptothrix rubra Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 63; Cladothrix rubra Mace, Traite Pratique de Bact., 4th ed., 1901, 1097; not Actinomyces ruber Krainsky, Cent. f. Bakt., II Abt., U, 1914, 662; Sanfelice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 355; Nocardia rubra Chalmers and Christopherson, Ann. Trop. :\Ied. and Parasit., 10, 1916, 265; Discomyces ruber Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 981.) From sputum. Some authors consider the following sj'nonyraous with this organism: Streptothrix rnineacea {Acti- nomyces mineaceus Lachner-Sandoval , Ueber Strahlenpilze, 1898, 66). Actinomyces rubidaureus Lachner- Sandoval. {Cladothrix mordore, Thiry, Arch. Physiol. Norm, et Path., 9, 1897, 283; Lachner-Sandoval, Ueber Strah- lenpilze, Inaug. Diss., Strassburg, 1898, 66; Actinomyces mordore, Thirj', These, Nancy, 1900, 82; Nocardia mordore, Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 265; Xocardia thiryei de Mello and Pais, Arq. Hig. Pat. E.xot., 6, 1918, 193; Dis- comyces thiryi Brumpt, Precis de Para- sitol., Paris, 3rd ed., 1922, 981; Oospora mordore, Sartory, Champ. Paras. Homme et Anim., 1923,824; Actinomyces thiryi Sartory and Bailly, Mycoses Pulmonaires, 1923, 252.) From a case showing anginous exudate with edema. Actinomyces salvati Langeron. (Lang- eron, Bull. Soc. Path. Exot., 15, 192?, 526; Fontoynont and Salvat, ibid., 596.) From generalized nodular lesions in the ^Madagascar rat. Actinomyces sarloryi Dodge. (Oo- spora pulmonalis var. acido-resistant, Sartory, Arch. Med. Pharm. .Milit., 70, 1916, 605; Dodge, Medical Mycology, St. Louis, 1935, 756.) From a patient showing symptoms of pulmonarj' tuber- culosis. Actinomyces septicus Mac Neal and Blevins. (Jour. Bact., 49, 1945, 605.) From human endocarditis. Actinomyces serratus Dodge. {Acti- nomyces asteroides var. serratus Sartory, Meyer and Meyer, Ann. Inst. Past., 44, 1930, 298; also see Compt. rend. Acad. Sci. Paris, 188, 1929, 745; Dodge, Medical Mycology, St. Louis, 1935, 745.) From a case of actinomycosis of bones with yellow grains. Actinomyces sommeri Greco. {Strepto- thrix hiadurae Greco, Primer Caso de 918 MANUAL OF DETERMINATIVE BACTERIOLOGY Pi^ de Madura o Micetoma en la Repii- blica Argentina, Thesis, Buenos Aires, 1904; Streptothrix inicetomas argentinac a, Greco, in Durante, Segunda Observa- cion de Pid de Madura o Micetoma en la Republica Argentina, Thesis, Buenos Aires, 1911; Greco, Origine des Tumeurs, 1916, 726; Nocardia micetomae-argentinae Durant, 1911, quoted from Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 985; Oospora sotnmeri Sartory, Champ. Paras. Homme et Anim., 1923, 783.) From a case of mycetoma pedis in Argen- tina. Actinomyces tossicus Dodge. (Acti- nomyces albus var. tossica Rossi, Ann. Ig. Sperim., 9, 1905, 693; Oospora alba var. toxique, Sartory, Champ. Paras. Homme et Anim., 1923, 829; Dodge, Medical Mycology, St. Louis, 1935, 719.) From tumors in the abdominal cavities of domestic fowl . Actinomyces urcthridis Brumpt. (Pre- cis de Parasitol., Paris, 4th ed., 1927, 1206.) Isolated by Rocek in 1920 from cases of prostatitis. Actinomyces variabilis Cohn. (Cent, f. Bakt., I Abt., Orig., 70, 1913, 301.) From pus in the bladder in a case of cystitis and from the prostate. Asleroidcs pscudocarneus Puntoni and Leonardi. (Boll, e Atti d. R. Accad. Med. di Roma, 61, 1935, 93.) Bacillus berestnewi Lcpeschkin. (Cent. f. Bakt., II Abt., 12, 1904, 641 and 13, 1904, 13.) From sputum of a pneumonia patient. Bacillus (Micrococcus?) havaniensis Sternberg. (Sternberg, Manual of Bact., 1893, 718; Bacterium havaniensis Ches- ter, Ann. Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 116; Serratia havaniensis Bergey et al., Manual, 1st ed., 1923, 95.) From human intestinal canal. Cladothrix matruchoti Mendel. (Men- del, Compt. rend. Soc. Biol., 82, 1919, 583; Nocardia matruchoti Pettit, 1921, quoted from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51; Oospora matruchoti, quoted from Nan- nizzi, idem; Actinomyces matruchoti Nan- nizzi, idem.) From the roots of a decay- ing tooth with tumefaction. Species dubia. Cohnistrepothrix misri Carpano. (Riv. di Parassitologia, n. 2, p. 107; cjuoted from Boll. d. Sez. Ital. d. Soc. Internat. d. Microbiol., 10, 1938, 62.) From human dermatosis in Egypt. Discomyces berardijiisi Brumpt. (Streptothrix sp. de Berardinis, Ann. Ottalmol. Lavori Clin. Oculist. Napoli, 33, 1904, 914; Actinomyces de Berardinis, Namyslowski, Cent. f. Bakt., I Abt., Orig., 62, 1912, 566; Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 977; Actinomyces bernardinisi Brumpt, ibid., 4th ed., 1927, 1189; Nocardia berardinisi Vuillemin, Encyclopedic Mycologique, Paris, 2, 1931, 126.) From a case of keratitis. Discomyces brasiliensis Lindenberg. (Lindenberg, Rev. med. de S. Paolo, 1909, No. 18, and Arch, de Parasitol., 13, 1909, 265; Nocardia brasiliensis Pino}^ Bull. Inst. Past., Paris, 11, 1913, 936; Streptothrix b7-asilie7isis Greco, Origine des Tumeurs, 1916, 724; Oospora brasiliensis Sartory, Champ. Paras. Homme et Anim., 1923, 786; Actinomyces brasiliensis Gomes, Ann. Palistas Med. Cirurg., 14, 1923, 150.) From a case of mycetoma of the leg. According to Pinoy and others, idential with Nocardia asteroides. Discomyces congolensis Baerts. (Baerts, Bull. Med. Katanga, 2, 1925, 67; Acti- nomyces congolensis Brumpt, Precis de Parasitol., Paris, 4th ed., 1927, 1206.) From lesions in a case of actinomycosis from the Belgian Congo. Discotnyces dispar (Vidal) Brumpt. (Microsporon dispar Vidal, 1882, Micro- sporon anemoeon Vidal, 1882 and Sporo- trichum dispar, quoted from Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 995 and from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 25; Brumpt, idem; Actinomyces dispar Brumi)t, ibid., 4th ed., 1927, 1206.) FAMILY ACTIXOMYCETACEAE 919 From a case of pityriasis. Species dubia. Discomyces mexicanus Boyd and Crutchfield. (Boyd and Crutchfield, Amer. Jour. Trop. Med., 1, 1921, 268; Actinomyces mexicanus Brumpt, Precis de ParasitoL, Paris, 4th ed., 1927, 1192; Nocardia mexicana Ota, Jap. Jour. Derm. Urol., 28, 1928.) From a mycetoma of the foot. Discomyces minutissimus (Burchardt) Brumpt. (Microsporon yjiinniissimum Burchardt, in Uhle and VVagn. Pat. Gen., 1859; Trichotheciiun sp. J. Neumann, 1868; Microsporon gracile Balzer, Ann. Derm. Syphiligr., II, 4, 1883, 681; Sporo- trichum ?ninuiissi77ium Saccardo, Sylloge Fungorum, 4, 1886, 100; Microsporoides minutissimus Xeveu-Lemaire, Precis ParasitoL, 1906; Brumpt, Precis de ParasitoL, 1st ed., 1910, 863; Oospora minutissima Ridet, Oospora et Oosporo- ses, 1911, 68; Nocardia minutissima Verdun, Precis parasitoL, 1912; Acti- nomyces minutissimus Brumpt, ibid., 4th ed., 1927, 1199.) Reported as the etiological agent of erythrasma. Mycobacterium alluvialum Bergej' et al. (Kersten, Cent. f. Bakt., I Abt., Orig., 51, 1909, 494; Bergey et aL, Manual, 1st ed., 1923, 379.) From soil. Mycobacterium convolutum Gray and Thornton. (Gray and Thornton, Cent, f. Bakt., II Abt., 73, 1928, 87; Actinomy- ces convolutus Bergey et al., Manual, 3rd ed., 1930, 473.) From soil. Resembles Nocardia opaca. Nocardia arborescens (Edington) Trevisan. {Bacillus arborescens Eding- ton, Brit. Med. Jour., June 11, 1887, 1262; Trevisan, I generi e le specie delle Batteriacee, 1889, 9; Actinomyces arbores- cens Gasperini, Cent. f. Bakt., 15, 1894, 684.) From human skin in cases of scarlatina. Nocardia bahiensis da Silva. (Da Silva, Brasil Med., 33, 1919, 81 and Mem. Inst. Butantan, 1, 1918-1919, 187, Dis- comyces bahiensis Neveu-Lemaire, Precis de ParasitoL Hmn., oth ed., 1921, 44; Oospora bahiensis Sartory, Champ. Paras. Homme et Anim., 1923, 784; Actinomyces bahiensis Brumpt, Precis de ParasitoL, 4th ed., 1927, 1195.) From an actinomycotic mycetoma in Brazil. Nocardia berestneffi Clialmers and Christopherson. (Streptothrix cas I, II, Berestneff, Inaug. Diss., Moscow, 1897; Chalmers and Christopherson, Ann. Trop. ]Med. and ParasitoL, 10, 1916, 263; Discomyces berestneffi. Brumpt, Precis de ParasitoL, Paris, 3rd ed., 1922, 992; Actinomyces berestneffl Sartory and Bailly, Mycoses pulmonaires, 1923, 256.) From a case of pulmonary pseudotuber- culosis. Nocardia bicolor (Trolldenier) de Mello and Fernandes. {Actinomyces bicolor Trolldenier, Ztschr. f. Tiermedizin, 7, 1903, 81; de Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 106.) Found in cerebromeningitis, bronchitis and lymphadenitis in a dog. Nocardia convoluta Chalmers and Christopherson. (Chalmers and Chris- topherson, Ann. Trop. Med. and Parasit., 10, 1916, 257; Discomyces convolutus Xeveu-Lemaire, Precis ParasitoL Hum., 5th ed., 1921, 44; Oospora convoluta Sartory, Champ. Paras. Homme et Anim., 1923, 769; Actinomyces convolutus Brumpt, Precis de ParasitoL, Paris, 4th ed., 1927, 1195.) From a yellow grain mycetoma of the foot in the Sudan. Nocardia cylindracea de Korte. (De Korte, Ann. Trop. Med. Parasit., 11, 1918, 205; Discomyces cylindraceus Neveu-Lemaire, Precis ParasitoL Hum., oth ed., 1921, 44; Oospora cylindracea Sartor}', Champ. Paras. Homme et Anim., 1923, 774; Actinomyces cylindra- ceus Brumpt, Precis de Parasitologic, Paris, 4th ed., 1927, 1206.) From an infection of the outer ear, resembling mycetoma. Nocardia enteritidis (Pottien) Castel- lan! and Chalmers. {Streptothrix enteri- tidis Pottien, 1902, according to San- felice, Cent. f. Bakt., I Abt., Orig., 36, 920 ^L\NUAL OF DETERMINATIVE BACTERIOLOGY 1904, 357; Bacillus enteriiidis Pottien, 1902, according to Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 25; not Bacillus cnteritidis Gaertner, Corre- spond, d. Allgemein. Artz. Vereins Thuringen, 17, 1888, 573; Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 819 ; Oospora enteritidis, quoted from Nan- nizzi, loc. cit.; Discomyces enteriiidis Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 980; Actinomyces enteritidis Brumpt, ibid., 4th ed., 1927, 1191.) From a case of enteritis. Nocardia eqiti Chahners and Christo- pherson. (Strcptothrix sp. Dean, Trans. Path. Soc. London, 5/, 1900, 26; Chalmers and Christ opherson, Ann. Trop. Med. Parasit., 10, 1916, 263; Discomcyes equi Brumpt, Precis de Parasitologie, 3rd ed., 1922, 992; vldmomyyces equi Sartory and Bailly, Mycoses Pulmonaires, 1923, 256.) From an abscess on the jaw of a horse. Nocardia crythrea (Foulerton) Chal- mers and Christopherson. (Strcpto- thrix crythrea Foulerton, 1910; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268.) Nocardia Joulertoni Chalmers and Christopherson. {Strcptothrix hominis Foulerton and Jones, Trans. Path. Soc. London, 53, 1902, 56; Strcptothrix hotninis I Foulerton, in Allbutt and IloUeston, Syst. of :\Iedicine, 2, 1906, 302; Nocardia hominis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 819; Oospora hominis Ridet, 1911, according to Castel- lani and Chalmers, idem; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 263; Discomyces jouler- toni Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 993; Actinomyces Joulertoni Brumpt, ibid., 4th ed., 1927, 1204.) From abscesses of the chest and from sputum. Nocardia genesii Froes. (Froes, Do mycetoma i)edis no Brasil 50, 1930; Actinomyces genesii Dodge, Medical Mycology, St. Louis, 1935, 761.) From a case of Genesio Salles from Bahia. Nocardia jollyi Vuillemin. (Vuille- min, in Jolly, Rev. Med. de I'Est, 48 1920, 42; Discomyces jollyi Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 84; Oospora jollyi Sartory, Champ. Paras. Homme et Anim., 1923, 770; Actinomyces jollyi Brumpt, ibid., 4th ed., 1927, 1196.) From a case clinically resembling bubonic plague. Nocardia lasserei Verdun. {Nocardia sp. Lasserre, These Toulouse, 1904; Verdun, Precis Parasitol., 1912; Dis- comyces lasserrei Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 43; Oospora lasserei Sartory, Champ. Paras. Homme et Anim., 1923, 798; .<4c/mow?/ces lasserei Brumpt, Precis de Parasitol., 4th ed., 1927, 1206.) From an ulcerative lesion on the pharynx and upper lip. Nocardia lingualis (Eisenberg) Chal- mers and Christopherson . (Zungenbelag- Vibrio, Weibel, Cent. f. Bakt., 4, 1888, 227; Vibrio lingualis Eisenberg, Bakt. Diag., 3 Aufl., 1891, 212; Spirillum liy^guae Sternberg, Man. of Bact., 1893, 697; Spirosoma lingualis Migula, in Engler and Prantl, Die natiirl. Pflan- zenfam., 1, la, 1895, 31; Strcptothrix lingualis Bajardi, Cent. f. Bakt., I Abt., Orig., 35, 1904, 129; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 265; not Nocardia lingualis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 819; Dis- comyces lingualis Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 980; Actinomyces lingualis Sartory and Bailly, Mycoses Pulmonaires, 1923, 252.) From a deposit on the tongue. Jensen (Proc. Linn. Soc. New So. Wales, 59, 1934, 43) regards this species as closely related to Coryncbacteriuin filatnentosum. Also see page 205. Nocardia londinensis (sic) Chalmers iind Christopherson. {Strcptothrix hominis II, Foulerton, Lancet, 1, 1900, 970; Chalmers and Christopherson, Ann. Troj). Med. and Parasit., 10, 1916, 256; Discomyces londinensis Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 993; Actinonujccs londinc7isis lirumpt, ibid., FAMILY ACTINOMYCETACEAE 921 4th ed., 1927, 1204.) From cases of human actinomycosis. Nocardia macrodipodidanun Fox. (Fox, Disease in Captive Wild Mammals and Birds, 1923, 570; Actinomyces macro- dipodidarum Dodge, INIedical Mycology, St. Louis, 1935, 747.) Found in lumpy jaw with septicemia and gasteroenteritis of kangaroos. Nocardia nicollci Delanoe. (Delanoe, Arch. Inst. Past. Tunis, 17, 1928, 257; Actinomyces nicollci Nannizzi, in Pol- lacci, Tratt. Micopat. Umana, 4, 1934, 36.) From a mycetoma of the thigh. Nocardia nigra (Castellani) Castellani and Chalmers. {Streptothrix nigra Cas- tellani, 1913, according to Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 1062; presumably not Strepto- thrix nigra Rossi Doria, Ann. 1st. Ig. sper. Univ. Roma, 1, 1891, 399; Castellani and Chalmers, idem.) From black gran- ules in a case of tonsillar nocardiomyco- sis. Usuall}^ forms black colonics on agar. Nocardia panginensis de Mello and Fernandes. (De Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 130; Actinomyces panginensis Dodge, Medical Mycology, St. Louis, 1935, 718.) From a dermatosis. Nocardia pijperi Castellani and Chalmers. (Nocardia sp. Pijper, Folia Microbiol., 5, 1917, 50 and Med. Jour. So. Africa, 12, 1917, 141; Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 1060; Discomyces pijperi Neveu- Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 44; Actinomyces pijperi Sartory and Bailly, Mycoses pulmonaires, 1923, 256.) From sputum in a case of chronic bronchitis. Nocardia pinoyi de Mello and Fernan- des. (De Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 130; Acti- nomyces pinoyi Dodge, Medical My- cology, St. Louis, 1935, 723.) From a case of erythrasma. Nocardia ponceti Verdun. (Verdun, Precis Parasitol., 1912, Discomyces pon- ceti Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 43; Oospora ponceti Sartory, Champ. Paras. Homme et Anim., 1923, 778; Actinomyces ponceti Brumpt, Precis de Parasit., Paris, 4th ed., 1927, 1205.) Isolated by Moorhof, Dor and Poncet from a muscular pseudo- actinomycosis. This species may be synonymous with Nocardia krausei (Chester). Nocardia putridogenes (Vezspremi) de ^Niello and Pais. (Cladothrix putrido- genes Vezspremi, Cent. f. Bakt., I Abt., U, 1907, 408 and 515; de :Mello and Pais, Arq. Hig. Pat. E.xot., 6, 1918, 187, Oo- spora putridogenes Sartory, Champ. Paras. Homme et Anim., 1923, 823; Actinomyces putridogenes Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 41 .) From greenish pus from gingi- val ulcers and abscess of the jaw. Nocardia repens (Eklund) Vuillemin. [Epidermophyton sp. Lang, 1879; Lepo- colla repens Eklund, 1883 ; Achorion repens Gueguen, 1904; Epidermophyton repens; all quoted from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934; Vuille- min, Encyclopedie Mycologique, Paris, 2, 1931, 124; Actinomyces repens Nan- nizzi, idem.) From the skin in cases of psoriasis. Nocardia rivierei Verdun. (Strepto- thrix sp. Riviere, Cong. Frang. INWd. Bordeaux 1895, 2, 1896, 1003; Actinomyces Sabrazes et Rivieri, Berestnev, Inaug. Diss., Moscow, 1897; Verdun, Pi-^cis Parasitol., 1912; Discomyces rivierei Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 43; Oospora rivierei Sartory, Champ. Paras. Homme et Anim., 1923, 792; Actinomyces rivierei Brumpt, Prdcis de Parasitol., 4th ed., 1927, 1201.) From sputum and pus in a bronchopleuropul- monary infection followed by multiple abscesses. Nocardia sanfelicei Redaelli . (Strepto- thrix acido-resistente, Sanfelice, Boll. 1st. Sieroterapico Milanese, 2, 1922, 327; Redaelli, 1st. Sieroterapico Milan- ese, 7, 1928, 75 and 121; Actinomyces 922 MANUAL OF DETERMINATIVE BACTERIOLOGY sanfclicei Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51.) From fatal lesions in a rat. Nocardia splenica Gibson. (Gibson, 1930, quoted from Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 50; Actinomyces splenica Nannizzi, idem.) From a case of splenomegalia. Nocardia tenuis Castellani. (Castel- lani, Brit. Jour. Derm. Syphilis, 23, 1911, 341; Discomyces tenuis Castellani, Proc. Roy. Soc. Med., 6, Derm., 1912, 23; Cohnistreptothrtx tenuis Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 273; Actinomyces tenuis Dodge, Medical Mycology, St. Louis, 1935, 715.) From cases of tricho- mycosis flava. Nocardia valvulae Chalmers and Christopherson. (Streptothrix valvulae destruens bovis Luginger, Monats. f. prakt. Tierheilk., 15, 1904, 289; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 263; Oospora valv^dae destruens bovis Sartory, Champ. Paras. Homme et Anim., 1923, 788; Actinomyces valvularis Ford, Textb. of Bact., 1927, 211; Actinomyces valvulae Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51.) From endocarditis in cattle. Oospora anuerobies Sartory. {Acti- nomyces sp. Butterfield, Jour. Inf. Dis., 2, 1905, 421; Sartory, Champ. Parasit. Homme et Anim., 1923, 830; Actinomyces anaerobies Plaut, quoted from Dodge, Medical Mycology, St. Louis, 1935, 717.) From pus from human lung. Oospora bronchialis Sartory and Levas- seur. (Sartory and Levasseur, 1914, quoted from Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 43; Actinomyces bronchialis Sartor}^, Bull. Sci. Pharm., 23, 1916, 12; Discomyces bronchialis Neveu-Lemaire, idem.) From sputum in a case of pulmonary oosporosis. Oospora buccalis Roger, Bory and Sar- tory. (Roger, Bory and Sartory, Bull. Mem. Soc. Med. Hop. Paris, 27, 1909, 319 and Compt. rend. Soc. Biol., 66, 1909, 301; Discomyces buccalis Brumpt, Precis de Parasitol., Paris, 1st ed., 1910, 861, Nocardia buccalis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 819; Actinomyces buccalis Sartory and Bailly, Mycoses pulmonaires, 1923, 256.) From a case of creamy stomatitis with tonsillar abscess. Oospora catarrhalis Sartory and Bailly. (Sartory and Bailly, These Univ. Stras- bourg Fac. Pharm., 4, 1921, 57; Dis- comyces catarrhalis Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 984; Actinomyces catarrhalis Brumpt, ibid., 4th ed., 1927, 1195.) From sputum in a case of pulmonary oosporosis. Oospora hominis Ridet. {Streptothrix hominis IV Foulerton, 1906; Streptothrix hominis III Foulerton, 1910; Ridet, 1911, according to Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 9; Nocardia appendicis Chalmers and Christopher- son, Ann. Trop. Med. and Parasit., 10, 1916, 256; Discomyces appendicis Brumpt, Precis de Parasitol., 3rd ed., 1922, 977; Actinomyces appendicis Brumpt, ibid., 4th ed., 1927, 1189.) From a case of appendicitis and an iliac abscess. Oospora lingualis Gu^guen. (Gudguen, Compt. rend. Soc. Biol., 64, 1908, 852; Discomyces lingualis Brumpt, Precis de Parasitol., Paris, 1st ed., 1910, 865 and 2nd ed., 1913, 976; not Discomyces lingua- lis Brumpt, ibid., 3rd ed., 1922, 980; Nocardia lingualis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 819; not Nocardia lingualis Chal- mers and Christopherson, Ann. Trop. Med. and Parasitol., 10, 1916, 265; Discomyces guegueni Brumpt, ibid., 3rd ed., 1922, 980; Actinomyces guegueni Brumpt, ibid., 4th ed., 1927, 1191, No- cardia guegueni Ota, Jap. Jour. Derm. Urol., 28, 1928.) From cases of lingua nigra. Oospora pulmonalis Roger, Sartory and Bory. (Roger, Sartory and Bory, Compt. rend. Soc. Biol., 66, 1909, 150; Discomyces pidmonalis Brumpt, Precis FAMILY ACTINOMYCETACEAE 923 de Parasitologic, 1st ed., 1910, 860; Nocardia pulmonalis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 817; Actinomyces pulmonalis Sar- tory and Bailly, Mycoses Pulmonaires, 1923, 256.) From the sputum of a patient with pulmonary mycosis. Oospora pulmonalis var. chromogena Sartory. (Sartory, 1913; Actinomyces pulmonalis var. chromogenus Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 39.) From sputum of a patient suspected of having puhnonary tubercu- losis. Proaclinomyces alhus Krassilnikov. (Bull. Acad. Sci., U. S. S. R., No. 1, 1938, 139.) Cells at first produce my- celium with frequent branching varying in diameter from 0.6 to 1.0 micron. Breaks up after 2 or 3 days into rods and sometimes later into cocci. Multiply by fission, cross -wall formation and budding. Does not form spores; cells are Gram-positive and are not acid-fast. Colorless growth. Colonies vary in the different strains, somewhat rough, folded, shiny or dull, of a dough-like consistency, Krassilnikov listed several strains of this organism, including Proactinomyces oli- gocarbophilus. Proactinomyces uquosus Turfitt. (Jour. Bact.. 47, 1944, 490.) From soil. Decomposes cholesterol. Proaclinomyces cyaneus (Beijerinck) Krassilnikov. (Actinococcus cyaneus Beijerinck, Folia microbiol., Delft, 3, 1914, 196; Krassilnikov, loc. cil.) Blue pigment produced on synthetic media. Cells are i-od-shaped. 0.7 to 0.8 by 3 to 7 microns. Branching cell material on potato multiplies by means of bud forma- tion, by fission, and cross-wall formation; no true spores formed. P roactinomyces cyaneus-antibiolicus Cause. (Jour. Bact., 51, 1946, 049.) From soil. Produces litmocidin, a new antibiotic. Proactinomyces moormani Franklin. (Ann. Intern. Med., 13, 1940, 1205.) From the pus of multiple molar abscesses in a dental patient. P roacli7iomyces paraguayensis Al- meida. (Mycopath., 2, 1940, 201.) From a thoracic mycetoma with heavy, dark grains affecting a Canadian patient living in the Paraguayan Chaco. Sabouraud's glucose agar: Pseudomem- branous colony with raised, dark center surrounded by a white band, progres- sively increasing in size, and then by a light chocolate area. Proactinomyces reslrictus Turfitt. (Jour. Bact., 47, 1944, 491.) From soil. Decomposes cholesterol. Proactinomyces sp. Helzer. Found in sputum of tuberculous patient. Patho- genic for guinea pigs and rabbits. Streptothrix buccalis Goadby. (M}'- cology of the Mouth, London, 1903, 200.) From the mouth in cases of pyorrhoea. Chalmers and Christopherson (Ann. Trop. Med. and Parasit., 10, 1916, 234) regard this as a synonym of Nocardia liquej aciens . Streptothrix flava Chester. {Acti- nomyces sp. Bruns, Cent. f. Bakt., 26, 1899, 11; Chester, Man. Determ. Bact., 1901, 362; Nocardia hruni Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 256; Streptothrix homi- nis Bruns, according to Chalmers and Christopherson, idem; Discomyces hruni Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 992; Actinomyces hnrni Brumpt, ibid., 4th ed., 1927, 1204; Acti- nomyces flavus Dodge, Medical My- cology, St. Louis, 1935, 752.) From pus from a case of actinomycosis of the ab- dominal wall. Streptothrix fusca Karwacki. (Kar- wacki, Compt. rend. Soc. Biol., 66, 1911, 180; not Streptothrix fusca Corda, Prachtflora europaischer Schimmel- bildungen, Leipzig and Dresden, 1839, 27; Nocardia fusca Castellani and Chal- mers, Man. Trop. Med., 2nd ed., 1913, 818; Discomyces fuscus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 993; Oospora fusca Sartory, Champ. Paras. 924 MANUAL OF DETERMINATIVE BACTERIOLOGY Homme et Anim., 1923, 809; Actinomyces fuscus Sartory and Bailly, Mycoses pulmonaires, 1923, 256; not Actinomyces fusca Sohngen and Fol, Cent. f. Bakt., II Abt., 40, 1914, 87.) From the sputum of a tuberculosis patient. Streptothrix litieola Foulerton and Jones. (Foulerton and Jones, Trans. Path. Soc. London, 53, 1902, 75; also see Foulerton, Lancet, 1, 1905, 1200 and 1, 1906, 970; Nocardia lutcola Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818; Discomyccs luteolus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 981; Oospora luteola Sartory, Champ. Paras. Homme et Anim., 1923, 812; Aclinoinyces luteolus Ford, Textb. of Bact., 1927, 213.) From a lung infection, from a case of conjunc- tivitis, and from a dental abscess. Streptothrix viadurae Solari. (Solari, Semana Med., 2^, 1917, 573; not Strepto- thrix madurae Vincent, Ann. Inst. Past., 8, 1894, 129; not Streptothrix madurae Koch and Stutzger, Ztschr. f. Hyg., 69, 1911, 17.) From Madura foot. See Actinomyces avadi. Streptothrix muris-ratti Schottmuller. (Schottmuller, Derm. Wochnschr., 58, 1914, 77; Nocardia 77uiris de Mello and Pais, Arq. Hig. Pat. Exot., 6, 1918, 183; Actinomyces muris-ratti Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 51.) From sodoku or rat-bite fever. Streptothrix tarozzii Miescher. {Acti- nomyces albiis Tarozzi, Archivio Sci. Med., 33, 1909, 553; Miescher, Arch. Derm. Syphilis, 124, 1917, 297; Actinomy- ces tarozzii Dodge, iVIedical Mycology, St. Louis, 1935, 735.) From a case of Madura foot. Streptothrix verrucosa (Adler) Miescher. (Actinomyces verrucosus Adler, 1904, see Xannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 46; Miescher, Arch. Derm. Syphilis, 124, 1917, 314.) From mycetoma pedis. FAMILY ACTIXOMYCETACEAE 925 Genus II. Actinomyces Harz. (Harz, in Bollinger, Centbl. f. med. Wissensch., 15, 1877, 485; Harz, Jahresber. d. Miinch. Thierarzneischule for 1877-78, 1879, 125; not Actinomyce Meyen, Linnaea, 2, 1827, 442; Discomyces Rivolta, Clinica Veter., Milano, /, 1878, 208; Actinocladothrix Afanasiev, St. Petersb. med. Wchnschr., 4, 1887, 323; Micromyces Gruber, Cent. f. Bakt., 10, 1891, 648; not Microtmjces Dangeard, Le Botaniste, 1, 1888, 55; Actino- bacierium Haas, Cent. f. Bakt., I Abt., Orig., 40, 1906, 180; Carieria and Carterii Musgrave, Clegg and Polk, Philippine Jour. Sci., Ser. B, Med. Sci., 3, 1908, 470; Cohnistrepiothrix Pinoy (in part), 1911, see Pinoy, Bull. Inst. Past., 11, 1913, 929; Anaeromyces Castellani. Douglas and Thomson, Jour. Trop. Med. Hyg., 24, 1921, 149; Brevistreptothrix Lignieres, Ann. Parasit. Hum. Comp., 2, 1924, 1.) True mycelium produced. The vegetative mycelium fragments into elements of irregular size and may exhibit angular branching. Xo conidia produced. Not acid- fast. Anaerobic to microaerophilic. Pathogenic for man and animals. The type species is Actinomyces bovis Harz. Key to the species of geinis Actinomyces. I. Colonies soft, smooth, uniform, not adherent to medium. Xo aerial hyphae. 1. Actinomyces bovis. II. Colonies tougher in texture and warted in appearance, adherent to medium. Scanty aerial growth of hyphae. 2. Actinomyces israeli. 1. Actinomyces bovis Harz. (Harz, in Bollinger, Cent. f. med. Wissensch., 15, 1877, 485; Jahrb. d. Miinch. Thier- arzeneischule, 1877, 781 ; Discomyces bovis Rivolta, La clinica Veterinaria, 1, 1878, 169 or 208; Bacterium actinocladothrix Afanasiev, St. Petersburger Med. Wochnschr., 13, 1888, 84; Nocardia ac- tinomyces Trevisan, I generi e le specie delle Batteriacee, Milan, 1889, 9 ; Strepto- trix actinomyces Rossi Doria, Ann. d. 1st. d'Igi. Sper., Univ. di Roma, 1, 1891, 425; Cladothrix bovis Mace, Traite de Bact., 2nd ed., 1891, 666; Oospora bovis Sauvageau and Radais, Ann. Inst. Past., Paris, 6, 1892, 271; Actinomyces bovis sulphureus Gasperini, Cent. f. Bakt., 15, 1894, 684; Nocardia bovis Blanchard, in Bouchard, Traits de Path. G^n^rale, 2, 1896, 857; Actinomyces sulphureus Gasperini, Atti Soc. Tosc. Scienz. Nat., P. v., 11, 1896; Cladothrix actinomyces Mace, Traite de Bact., 3rd ed., 1897, 1038; Streptothrix actinomycotica Fouler- ton, Lancet, 2, 1899, 780; Streptothrix bovis communis Foulerton, Jour. Comp. Path, and Therap., 14, 1901, 50; Strepto- thrix bovis Chester, Man. Determ. Bact., 1901, 361, Streptothrix sulphurea Caminiti, Cent. f. Bakt., I Abt., Orig., 44, 1907, 197; Sphaerotilus bovis Engler, Syllabus der Pflanzenfam., 5 Aufl., 1907, 5; Nocardia sulphurea Vuillemin, Encj-clopedie Mj'cologique, Paris, 2, 1931, 129; Proactinomyces bovis Henrici, Biology of Bact., 2nd ed., 1939, 409.) From Latin bovis, of the ox. Synonyms previous to 1919 as given by Breed and Conn, Jour. Bact., 4, 1919, 596. Probable synonym : Brevistreptothrix spitzi Lignieres, Annales de Parasit., 2, 1924, 2 {Streptothrix spitzi Lignieres and Spitz, Cent. f. Bakt., I Abt., Orig., 35, 1904, 453; Actinobactcrium isreali var. spitzi Sampietro, Ann. Igiene, 18, 1908, 391; Discomyces spitzi Brumpt, Precis de Parasitol., Paris, 1st ed., 1910, 847; Actinomyces spitzi Lieske, Morphol. u. Biol. d. Strahlenpilze, Leipzig, 1921, 32; Oospora spitzi Sartory, Champ. Paras. Homme et Anim., 1923, 775). Found in mycosis of the upper jaw of oxen in Argentina. 926 MANUAL OF DETERMINATIVE BACTERIOLOGY Description from Erikson, Med. Res. Council, London, Special Report Ser. 240, 1940, 63 pp. No aerial hyphae. Radiate, sulfur- colored granules occur in the pus found in cases of actinomycosis. Large club- shaped hyphae are seen in morbid tissues. Gram-positive. Xon-motile. Not acid- fast. Mycelium : Undergoes fragmentation very rapidly, extensive branching is rare. Hyphae less than 1 micron in diameter. Colonies : Smoother and softer in con- sistency, and more uniform than in the following species. The colonies are not adherent to the medium and growth is scantier. Semi-solid media: E.xcellent growth, especially with paraffin seal. Gelatin : Occasionally scant, flaky growth. No liquefaction. Liquid media : Occasional turbidity with a light flocculent growth. Acid from glucose, sucrose and maltose. No acid from salicin and mannitol. Pigments : No soluble pigments pro- duced on protein media. No insoluble pigments produced by growth. Egg or serum media: No proteolytic action. Litmus milk : Becomes acid but usually no coagulation, no peptonization. Some- times no growth. No hemolysis in blood broth or blood agar. Serology: No cross agglutination be- tween five bovine strains and human strains of Actinomyces israeli. No cross reactions with representative aerobic strains. Optimum temperature 37°C. Anaerobic to microaerophilic. Bovine strains are more oxygen-tolerant on egg or serum media than strains of human origin belonging to the following species. As pointed out by Lignieres and Spitz (Bull. Soc. cent. Med. vet., 20, 1902, 487 and 546) and others, distinction should be made between the infections produced by Actinomyces bovis and those produced by the Gram-negative Actinobacillus now known as Actinobacillus lignieresi. These infections frequently occur in mixed form and are also frequently com- plicated by the presence of pyogenic cocci (Magnussen, Acta path. Microbiol. Scand., 5, 1928, 170; and others). Source : Originally found in lumpy jaw of cattle. Habitat : Frequently found in and about mouth of cattle and probably other animals. Lesions may also be produced in the liver, udder or other organs of cattle and hogs. Possibly also in human mouth (Naeslund, Acta path. Microbiol. Scand., 2, 1925, 110). This and the following species are sometimes regarded as being identical (see Emmons, Public Health Repts., U.S.P.H.S., 53, 1935, 1967; Rosebury, Bact. Rev., 8, 1944, 190; and others). 2. Actinomyces israeli (Kruse) Lachner-Sandoval. (Strahlenpilz, Wolff and Israel, Arch. f. path. Anat., 126, 1891, 11; Streptothrix israeli Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 56; Actinomyces israeli Lachner- Sandoval, Inaug. Diss., Strassburg, 1898, 64; Discomyces israeli Gedoelst, Les champignons parasites de I'homme et des animaux domestiques, 1902, 163; Actinobacterium israeli Sampietro, Ann. d'Ig. sperim., 18, 1908, 331 ; Cohnistrepto- thrix israeli Pinoy, Bull. Inst. Pasteur, Paris, 11, 1913, 931; Nocardia israeli Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 814; Anaeromyces bronchitica Castellani, Douglas and Thomson, Jour. Trop. Med. Hyg., 24, 1921, 149; Cohni streptothrix bronchitica Verdun and Mandoul, Precis Parasitol., 1924, 754; Brevistreptothrix israeli Lig- nieres, Annales de Parasit., 2, 1924, 2; Proactinomyces israeli Negroni, Compt. rend. Soc. Biol., Paris, 117, 1934, 1239; Corynebacterium israeli Haupt and Zeki, Cent. f. Bakt., I Abt., Orig., 130, 1933, [}o;Ons])ora israeli, quoted from Nanuizzi, FAMILY ACTIXOMYCETACEAE 927 Tratt. Micopat. Umaiia, 4, 1934, 53; Actinomyces wolff-israel Lentze, Cent. f. Bakt., I Abt., Orig., I4I, 1938, 21.) Named for Prof. Israel, one of the origi- nal isolators of this organism. Synonyms previous to 1919 essential Ij' as given by Breed and Conn, Jour. Bact., 4, 1919, 597. Description from Erikson, Med. Res. Council, London, Special Rept. Ser. 240, 1940, 63 pp. Erect aerial hyphae produced in an atmosphere of reduced oxygen tension. These hyphae are occasionally septate but no definite spores are formed. One micron or more in diameter. Large club-shaped forms are seen in morbid tissues. Gram-positive. Xon-motile. Not acid-fast. Substrate mycelium: Initially uni- cellular and the branches may extend into the medium in long filaments or may, more or less quickly, exhibit frag- mentation and characteristic angular branching. The latter resembles the similar phenomenon found in Corynehac- teriwn . Colonies : These exhibit a considerable degree of polymorphism but no stable variants have been established. Tougher in texture than those of Actinomyces bonis. Old colonies warted in appearance. Adherent to the medium. Gelatin: Occasionally scant, flaky growth. No liquefaction. Liquid media: L^sually clear. Acid from sugars: According to Slack (Jour. Bact., 43, 1941, 193-209) acid from glucose, maltose, mannitol, sucrose and lactose; according to Negroni and Bon- figlioli (Physics, 15, 1939, 159) acid from glucose, galactose, lactose, fructose, mal- tose, raffinose, sucrose and xylose. Pigments : No soluble pigments on protein media. No insoluble pigments produced by growth. Egg or serum media: No proteolytic action. Litmus milk : Becomes acid but usuallv does not clot. No peptonization. Fre- quently no growth. No hemolysis. Serology : No cross agglutination be- tween 12 human strains and bovine strains of Actinomyces. No cross reac- tions with representative aerobic strains. Optimum temperature 37°C. Anaerobic to microaerophilic. Source : From 2 cases of human ac- tinomycosis : (1) A retromaxillary tumor, (2) actinomycosis of lung and breast (Wolff and Isreal). Habitat : From human sources (mouth, tonsillar crypts, etc.). Also reported from various domestic animals such as dogs (Baudet, Ann. Parasit., 12, 1934, 296) and cats (Edington, Vet. Record., U, 1934, 311). Appendix: The following names have been applied to anaerobic or semi -anaero- bic species, with descriptions which do not permit clear separation from the above or from each other. Actinobacterium meyeri Pr6vot. (An- aerobe Streptothrix-Art, Meyer, Cent, f . Bakt., I Abt., Orig., 60, 1911, 75; Pr^vot, Ann. Inst. Past., 60, 1938, 303.) From fetid pus. Actinomyces discofoliatus Griiter. (Griiter, Ztschr. f. Augenheilk., 13, 19.33, 477; redescribed by Negroni, Mycopathologia, 1, 1938, 81.) From lachrj'mal concretion and human infec- tion. Actinomyces lanfranchii Sani. (Saui, 1916, quoted from Dodge, Medical Mj- cology, St. Louis, 1935, 731; Nocardia lanfranchii de Mello and Pais, Arq. Hig. Pat. Exot., 6, 1918, 178.) From glandu- lar and ganglionar actinom3'cosis of the ox. Regarded as a variety of Actinomy- ces bovis. Actinomyces thjoettae Dodge. (Cokni- slreptothrix sp. or Streptothrix sp. Thj0tta and Gundersen, Jour. Bact., 10, 1925, 1; Dodge, Medical INIycology, 1935, 713.) From the blood in a case of acute rheumatism. 928 MANUAL OP DETERMINATIVE BACTERIOLOGY Cohnistreptolhrix neschezadimenki (sic) Chalmers and Christopherson. (Eine Streptothrix, Xeschczadimenko, Cent. f. Bakt., I Abt., Orig., ^6, 1908, 573; Chalmers and Christopherson, Ann. Trop. Med., 10, 1916, 273; Actinomyces neschezadimenki Dodge, Medical My- cology, 1935, 712; Actinobacterium absces- sus Pr6vot, Ann. Inst. Past., 60, 1938, 303.) From a human infection. Discomyces carougeaui Gougerot. (Gougerot, Compt. rend. Soc. Biol., Paris, 67, 1909, 580; Nocardia carougeaui Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 817; Cohnistrepto- lhrix carogeanui (sic) Chalmers and Christopherson, Ann. Trop. Med., 10, 1916, 273; Streptothrix carougeaui Greco, Origine des Tumeurs, 1916, 724; Actino- myces carougeaui Brumpt, Precis de Parasitol., 4th ed., 1927, 1206.) From a human infection. Discomyces thibiergei Ravaut and Pinoy. (Ravaut and Pinoy, Ann. Derm, et Syph., 10, 1909, 417; Nocardia thi- biergei Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 817; Coh7ii- streptothrix thibiergei Pinoy, Bull. Inst. Past., 11, 1913, 938; Oospora thibiergei Sartory, Champignons Parasites de 1 'Homme et des Animaux, Fasc. 11, 1923, 792; Actinomyces thibiergei Greco, Ori- gine des Tumeurs, 1916, 723.) From a human infection. Streptothrix cuniculi Schmorl. (Schmorl, Deutsch. Ztschr. f. Thiermed., 17, 1891, 375; Actinomyces cuniculi Gasperini, Ann. 1st. Ig. sper. Univ. Roma, 2, 1892, 222; Cladothrix cuniculi Mac^, Traite de Bact., 6th ed., 2, 1913, 753; Cohnistreptothrix cuniculi Chalmers and Christopherson, Ann. Trop. Med., 10, 1916, 273; Nocardia cuniculi Froilani de Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 107; Oospora cumculi Sartory, Champignons parasites de 1 'Homme et des Animaux, Fasc. 11, 1923, 824.) From infections in rabbits. Gen- erally regarded as probably identical with Spherophorus necrophorus (Fliigge) Pr^- vot. See page 678. FAMILY STREPTOMYCETACEAE 929 FAMILY III. STREPTOMYCETACEAE WAKSMAN AND HENRICI.* (Jour. Bact., 46, 1943, 339.) Vegetative mycelium not fragmenting into bacillary or coccoid forms. Conidia borne on sporophores. Primarily soil forms, sometimes thermophilic in rotting manure. A few species are parasitic. Key to the genera oj family Streptomycetaceae. I. Conidia produced in aerial hyphae in chains. Genus I. Streptoynyces, p. 929. II. Conidia produced terminally and singly on short conidiophores . Genus II. Micromonospora, p. 978. Genus I. Streptomyces Waksman and Henrici. {Streptothrix Cohn, Beitr. zur Biol, der Pflanzen, /, Heft 3, 1875, 186; not Strep- tothrix Corda, Prachtfiora Europaescher Sohinmielbildung, 1839; Nocardia Wright, Jour. Med. Research, IS, 1905, 349; Nocardia Winslow et al.. Jour. Bact., 2, 1917, 554; not Nocardia Trevisan, I generi e le specie delle Batteriacee, 18S9, 9; Cohnistreptothrix (Group I) 0rskov, Investigations in the Morphology of the Ray Fungi. Copenhagen, 1923, 147; not Cohnistreptothrix Pinoy, in Li^gard and Lan- drieu. Bull. Soc. Opht., U, 1911, 253 and Bull. Inst. Past., 11, 1913, 929; Aerothrix Wollenweber (in part), Ber. deut. Bot. Gesel., 39, 1921, 26; Euactinomyces Langeron (in part), Nouv. Traits de Med., 4, 1922; Waksman and Henrici, Jour. Bact., 46, 1943, 339.) Organisms growing in the form of a much-branched mycelium with a typical aerial mycelium. Conidiospores formed in chains. Aerobic. Saprophytic soil forms, less commonly parasitic on plants or animals. This genus can be divided, on the basis of the structure of sporulating hyphae, into five groups : Group 1. Straight sporulating hyphae, monopodial branching, never producing regular spirals. Group 2. Spore-bearing hyphae arranged in clusters. Group 3. Spiral formation in aerial mycelium; long, open spirals. Group 4. Spiral formation in aerial mycelium; short, compact spirals. Group 5. Spore -bearing hyphae arranged on mycelium in whorls or tufts. The type species is Streptomyces albus (Rossi Doria emend. Krainsky) Waksman and Henrici. Key to the species of genus Streptomyces. I. Saprophytes; psychrophilic to mesophilic. A. Soluble pigment on organic media faint brown, golden-yellow, or blue; pigment may also be absent entirely. 1. Pigment absent, or faint brown pigment formed at first and later lost; aerial mycelium abundant, white. 1. Streptomyces albus. 2. Pigment blue or red, when present. The red (insoluble) phase occurs when the reaction is distinctly acid, the blue (soluble) phase when it is alkaline. 2. Streptomyces coelicolor. * Revised by Prof. S. A. Waksman, New Jersey Experiment Station, New Bruns- wick, New Jersey and Prof. A. T. Henrici, University of Minnesota, Minneapolis, Minnesota, May, 1943. 930 MANUAL OP DETERMINATIVE BACTERIOLOGY 3. Pigmeut at first green becoming brown; aerial mycelium usually absent. 3. Slreptomyces verne. 4. Pigment yellowish-green; growth on sjmthetic agar penetrating into medium, pink. 4. Streptomyces calif ornicus. 5. Pigment golden-yellow; growth on synthetic agar yellow, with yellow soluble pigment. 5. Streptomyces flaveolus. 6. Pigment brown (only on certain protein media, as gelatin, glucose broth). a. Grown on synthetic agar red to pink. Scant, white aerial mycelium. 6. Streptomyces bobiliae. aa. Growth on sj^nthetic agar colorless; aerial mycelium thin, rose-colored. 7. Streptomyces roseochromogenus. aaa. Growth on synthetic agar mouse-gray ; powdery aerial myce- lium. 8. Streptomyces griseolus. aaaa. Growth on synthetic agar white turning yellowish, aerial mycelium white. 9. Streptomyces eryihreus. B. Soluble yellow pigment on Ca-malate agar. 1. Proteolytic action strong in milk and gelatin. a. Yellow pigment formed. b. Cellulose decomposed; starch is hydrolyzed. 10. Streptomyces cellulosae. bb. Cellulose not decomposed. 11. Streptomyces parvus. 2. Proteolytic action weak. 12. Streptomyces malenconi. C. Soluble brown pigment formed on synthetic agar. 1. Yellowish-green pigment on potato. 13. Streptomyces diastaticus. 2. Red-brown pigment on potato plug. 14. Streptomyces fimicarius. D. Greenish-yellow soluble pigment formed; sulfur-yellow pigment on potato. 15. Streptomyces flavovirens. E. Soluble brown pigment formed in all media containing organic substances. 1. Pigment deep brown (chromogenic type). a. Pigment faint brown on organic media, becoming greenish- brown to black; reddish aerial mycelium on glucose agar. 16. Streptomyces olivochromogenus . aa. Aerial mycelium yellowish with gray margin; weak diastatic action. 17. Streptomyces diastatochromogenes. aaa. Aerial mycelium yellowish; diastatic action weak. 18. Streptomyces flavochromogenes. aaaa. Aerial mycelium gray ; sporophores in clusters; strongly anti- biotic. 19. Streptomyces antibioticus . FAMILY STREPTOMYCETACEAE 931 2. Growth and aerial mycelium green on synthetic agar. 20. Sireptomyces viridochromogenes . 3. Deep brown to black pigment on synthetic agar. a. Orange-red on potato; no aerial mycelium on synthetic agar; growing feebly. 21. Streptomyces purpeochromogenus . aa. Brown to black on potato; abundant cottony aerial mycelium on synthetic agar, b. Brown ring on milk culture; coagulated; peptonized. 22. Streptomyces phaeochromogenus. bbb. Black ring on milk; no coagulation ; peptonization doubt- ful. 23. Streptomyces aureus. c^ Red to rose-red pigment on glucose, maltose, and starch agar. 24. Streptomyces erytkrochromogenes . c^. Lavender-colored aerial mycelium. 25. Streptomyces lavendulae. c^. Growth on potato gray with black center. 2G. Streptomyces reticuli. c*. Growth on potato cream-colored, becoming pink to dark red. 27. Streptomyces rubrireticuli. c^. Growth on potato greenish-olive. d. Aerial mycelium straw-colored. 28. Streptomyces flavus. dd. Aerial mycelium chrome-orange. 29. Streptomyces ruber. F. No soluble pigment formed on gelatin or other media. 1. Proteolytic action strong in milk and gelatin. a^. Yellowish-green growth on starch with pinkish aerial mycelium. 30. Streptomyces citreus. a-. Golden-yellow growth, later becoming orange to red-brown, on synthetic media. 31. Streptomyces fulvissirjius. a'. Cream-colored growth on starch media. 32. Streptomyces gougeroti. a.*. Bluish-black color on synthetic media, with white aerial myce- lium. 33. Streptomyces violaceoniger . a^. Yellowish pigment on potato. b. Aerial mycelium thick, powdery, water-green; starch is hj-- drolyzed. 3-1. Streptomyces griseus. bb. Aerial mycelium white; starch weakly hydrol3^zed. 35. Streptomyces griseoflavus . a^. Greenish-black pigment on potato; aerial mycelium white. 36. Streptomyces alhido flavus. aJ . Reddish-brown pigment on potato; aerial mycelium white; starch is not h3'drolyzed. 37. Streptomyces poolensis. 932 MANUAL OF DETERMINATIVE BACTERIOLOGY a*. Gray to sulfur-yellow pigment on potato. b. Aerial mycelium mouse-gray to light drab; starch is hydrolyzed. 38. Streptomyces olivaceus. bb. Aerial mycelium yellowish-wiiite. 39. Streptomyces lieskei. bbb. No aerial mycelium; starch is hydrolyzed. 40. Streptomyces microflavus. a'. No soluble pigment on potato. 41. Streptomyces cacaoi. 2. Proteolytic action weak. a. Soluble pigment formed on synthetic agar, b. Pigment blue or blue-black. 42. Streptomyces novaecaesareae. bb. Pigment brown to black. 43. Streptomyces exfoliatus. 44. Streptomyces gelaticus. aa. No soluble pigment on synthetic agar. b^ Growth turning black; diastatic action strong. c. Growth on synthetic agar scant with abundant spirals in aerial mycelium. 45. Streptomyces rutgersensis. cc. No spirals on synthetic agar ; characteristic green- colored growth on protein-glycerol medium. 46. Streptomyces lipmanii. ccc. No spirals on synthetic agar; growth dark, almost black. 47. Streptomyces halstedii. b". Moist aerial mycelium, many spirals. 48. Streptomyces hygroscopicus. h^. Growth orange-colored. 49. Streptomyces fradiae. b*. Growth yellowish. 50. Streptomyces alboflavus. h^. Growth rose to red; aerial mycelium white. 51. Streptomyces albosporeus. b^. Growth cream-colored; aerial mycelium flaky. 52. Streptomyces flocculus. b'. Growth red; aerial mycelium black. c. Complete decomposition of cellulose; weakly diastatic. 53. Streptomyces melanosporeus . cc. Incomplete decomposition of cellulose; strongly diastatic. 54. Streptomyces melanocyclus. 3. No proteolytic action or very little. a. Acid-resistant strains. 55. Streptomyces acidophilus. aa. Non-acid-resistant. 56. Streptomyces rubescens. FAMILY STREPTOMYCETACEAE 933 II. Saprophytes; thermophilic. A. Starch hydrolyzed. Yellowish growiih on potato. 57. Streptomyces Ihermophilus . B. Starch not hydrolyzed. Abundant, dark-colored growth on potato. 58. Streptomyces thermofuscus . III. Plant parasites. A. Tyrosinase reaction positive ; aerial mycelium gray -white. 59. Streptomyces scabies. B. Tyrosinase reaction negative; attacks sweet potatoes. 60. Streptomyces ipomoea. IV. Isolated from animal tissues. In the animal body, hyphae often show clavate enlargements at the ends. A. Limited proteolytic action in gelatin, milk, coagulated egg-albumin or fibrin. 1. Color of vegetative growth golden-brown. 61. Streptomyces fordii. 2. Color of vegetative growth pink. a. Sparse white aerial mycelium. 62. Streptomyces africanns. aa. Formation on mycelium of bodies similar to Thermoactinomyces . 63. Streptomyces gallicus. aaa. Yellowish-pink growth on potato plug; scant white aerial mycelium. 64. Streptomyces pelletieri. 3. Color of vegetative growth white. 65. Streptomyces lister i. 4. Vegetative growth cream-colored, scant white aerial mycelium. 66. Streptomyces upcottii. 5. Growth very limited on various media, except on potato plug; no liquefaction of gelatin. 67. Streptomyces hortonensis . B. Strong proteolytic action in gelatin and milk. 1. Xo pigment produced. a'. No growth on potato plug. 68. Streptomyces gibsonii. a^. Moist, membranous growth on potato plug; diastase formed. 69. Streptomyces beddardii. 70. Streptomyces kimberi. a^. Extensive growth on potato media; white powdery aerial mycelium, a* Abundant growth on potato plug, becoming black; white-gray aerial mycelium; plug discolored. 71. Streptomyces somaliensis. a*. Pink-colored growth on some media. 72. Streptomyces panjae. a^. Profuse white aerial mycelium on most media, spiral formation. 73. Streptomyces willmorei. 1. Streptomyces albus (Rossi Doria (Streptotrix alba Rossi Doria, Ann. d'Ist. emend. Krainsky) Waksman and Henrici. d'Ig. sper. di. Univ. di Roma, 1, 1891, 934 MANUAL OF DETERMINATIVE BACTERIOLOGY 399; Cladothrix alba Mac6, Traitd Pra- tique Bact., 3rd ed., 1897; Actinomyces albus Krainsky, Cent. f. Bakt., II Abt., 41 , 1914, QQ2;Nocardia alba Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Waksman and Henrici, Jour. Bact., 46, 1943, 339.) From Latin albus, white. Additional synonyms as given by Baldacci (Mycopathologia, 2, 1940, 156) : Cladothrix dichotoma Mac^, not Cohn, 1886; Streptothrix foersteri Gasperini, not Cohn, 1890; Streptothrix No. 2 and 3, Almquist, 1890; Actinomyces saprophyti- cus Gasperini, 1892; Oospora doriae Sauvageau and Radais, 1892; Cladothrix liquefaciens Hesse, 1892 (according to Duch^) ; Cladothrix invulnerabilis Acosta and Grande Rossi, 1893; Actinomyces chromogenus Gasperini, 1894 {Streplo- trix nigra Rossi Doria, 1891); Strepto- thrix gedanensis I Scheele and Petruschky, 1897; Streptothrix gramine- arum Berestneff, 1898; Actinomyces ther- mophilis (Berestneff) Miehe, not Gilbert, 1898; Cladothrix odorifera Rullmann, 1898; Actinomyces chromogenes Gasperini j8 alba Lehmann and Neumann, 1899; Oospora sp. Bodin, 1899 (according to Duche) ; Oospora alpha Price-Jones, 1900 (according to Chalmers and Christopher- son) ; Streptothrix leucea Foulerton, 1902 (according to Chalmers and Christopher- son); Streptothrix Candida Petruschky, 1903; Streptothrix lathridii Petruschky, 1903; Streptothrix dassonvillei Brocq- Rousseau, 1907 (according to Duche) ; Streptothrix pyogenes Caminiti, 1907 (ac- cording to Chalmers and Christopher- son); Streptothrix sanninii Ciferri, 1922; Actinomyces almquisti Duche, 1934; Actinomyces gougeroti Duche, 1934. Doubtful synonyms : Oospora melchni- kowi Sauvageau and Radais, 1892; Oospora guignardi Sauvageau and Radais , 1892; Actinomyces albus Waksman and Curtis, 1919 ; Aclinomyces thermodiastal- icus Bergey, (1919) 1925. Varieties: Actinomyces albus var. acidus Xeukirch, 1902 (according to Nannizzi); Actino- myces albus var. ochroleucus Neukirch, 1902 (according to Wollenweber) ; Ac- tinomyces albus var. toxica Rossi, 1905; Actinomyces albus var. cretaceus (Kriiger) Wollenweber, 1920; Actino- myces albus var. a Ciferri, 1927. More complete information regarding these species will be found in the text or in the Appendix to Genus Streptomyces. The description of this species by Rossi Doria is incomplete. The charac- ters given below are taken from Krainsky {loc. cit.) with some supplementary in- formation from later authors. Other de- scriptions which may vary from this in certain details are given by Waksman and Curtis (Soil Sci., /, 1916, 117), Bergey et al. (Manual, 1st ed., 1923, 367), Duche (Les actinomj'ces du groupe albus, Paris, 1934, 257) and Baldacci {loc. cit.). Vegetative hyphae : Branched, 1 micron in diameter. Aerial mycelium: Abundant, white. Hyphae 1.3 to 1.7 microns in diameter with ellipsoidal spores (1 micron long) in coiled chains on lateral branches of the aerial hyphae. Gelatin: Liquefaction. Colonies gra}% no soluble pigment. Ca-malate agar : Colonies of medium size, the center onlj' is covered with a white aerial mycelium. Starch agar: Aerial mycelium white but covers the whole surface. Glucose agar : Gray aerial mycelium becoming brownish. Peptone and bouillon agar : No aerial mycelium but a chalkj^ white deposit forms on old colonies. Odor : Earthy or musty. Broth : Flaky growth on bottom with surface pellicle in old cultures. Potato: Colonies and aerial mycelium white. Carrots and other vegetables : Excel- lent growth (Duche). No growth on cellulose. No hydrolysis of starch. Activcl}^ proteolytic. Nitrites produced from nitrates. li'AMILY STREPTOMYCETACEAE 935 Milk: Peptonized after coagulation. Reaction becomes alkaline (Duch^) . Aerobic . Source : From air and soil (Rossi Doria) ; from garden soil (Krainsky) . Habitat: Dust, soil, grains and straw. Widely distributed. 2. Streptomyces coelicolor (Reiner- ^liiller) comh. nov. {Slreptothrix coelicolor Reiner-Mijller, Cent. f. Bakt., I Abt., Orig., 46, 1908, 197; Nocardia coelicolor Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271; Actinomyces coelicolor Lieske, Morphol. u. Biol. d. Strahlenpilze, Leipzig, 1921, 28.) From Latin caelum, sky and color. color. Regarded by the authors of this section as the same as Actinomijccs violaceus Waksman and Curtis, Soil Science, 1, 1916, 110 (Actinomyces violaceus-ruber Waksman and Curtis, ibid., 127; Actino- myces waksmanii Bergej^ et al., Manual, 3rd ed., 1930, 489) and Actinomyces tri- color Wollenweber, Arbeiten d. For- schungsinstitut fiir Kartoffelbau, 1920, 13. It is, however, pointed out by J. E. Conn (Jour. Bact., 46, 1943, 133) tJiat certain differences between the descrip- tions of Waksman and Curtis, and that of Miiller may correspond to actual chemical differences in the pigments produced; and that the organism of Waksman and Curtis may be a separate species. Description by Miiller except as noted. Morphology of Streptomyces coelicolor has not been fully described. According to Waksman and Curtis who described Actinomyces violaceus-ruber, this is as follows : Straight filaments with open, dextrorse spirals, breaking up into coni- dia. Conidia oval or rod -shaped, 0.7 to 1.0 by 0.8 to 1.5 microns. Gelatin: Good growth. Xo pigment formation. Liquefaction fairlj^ rapid, beginning in 4 to 7 days . Plain agar: Good growth. Pigment lacking or faint blue (Conn). Czapek agar (according to Waksman and Curtis concerning Actinomyces viola- ceus-ruber) : Thin, spreading, colorless at first, becoming red, then blue. Aerial mycelium thin, white, powdery, becom- ing mouse -gray. Asparagine agar (synthetic) : With glycerol as source of carbon, good growth, violet to deep blue, with pigment diffus- ing through medium; final H-ion con- centration about pH 7.0 to 8.0. With glucose as source of carbon, poorer growth, red, no diffusion of pigment; final H-ion concentration about pH 6.0 to 5.0 (Conn). Broth : Good growth. Cretaceous layer around edge. :VIilk: Xo change at 25°C (Conn). At 37°C, coagulation. Peptonization be- ginning in 3 to 5 da3-s. Potato : Strong pigment production, sometimes greenish-blue or violet, but usually sky-blue, diffusing through me- dium and coloring water at base of tube. Xitrites produced from nitrates. Blood agar : Hemolysis showing on 4th day. Miiller reports no acid from carbohy- drates on organic media. Conn, however, finds acid from glucose and lactose, and sometimes from sucrose and mannitol when grown on synthetic media. Pigment : The most striking charac- teristic of this organism is a litmus -like pigment usuallj- produced on potato or synthetic media, which is deep blue and water-soluble at alkaline reactions (be- yond pH 8.0), violet around neutralitj-, and red (insoluble in water) at about pH 6.0. Conn points out that the primary pigment has a spectrophotometric curve almost identical with that of azolitmin; but that there are undoubtedly other pig- ments produced, especially in the case of the strains believed to be typical of Actinomyces violaceus-ruber (as pre- viously pointed out by Waksman and Curtis). Good growth at room temperature and at 37°C. 936 MANUAL OF DETERMINATIVE BACTERIOLOGY Aerobic. Distinctive character : Litmus-like pig- ment. Source : Dust contamination on a potato slant. Habitat: Soil and plant surfaces. Very abundant. Note : Because of the numerous colors and shades shown by the pigment ac- cording to final H-ion concentration and other less understood factors, this species may have been described under various names. On the other hand, it is entirely possible, as pointed out by Conn {loc. cit.), that careful study of the pigments may show that more than one species is actually involved. 3. Streptomyces verne (Waksman and Curtis) comb. nov. (Actino^nyces verne Waksman and Curtis, Soil Science, 1, 1916, 120.) Derivation uncertain. Filaments with close branching of the hyphae. No conidia demonstrated. Gelatin stab: Small, cream-colored colonies. Rapid liquefaction. Synthetic agar: Abundant, spreading, wrinkled, elevated, glossy, yellowish growth, becoming brownish, lichenoid margin. Starch agar: Scant, brownish, re- stricted growth. Glucose agar: Abundant, much folded growth, center raised, gray with purplish tinge, entire. Plain agar : Small, grayish colonies with depressed center, becoming wrinkled. Glucose broth: Slightly flaky sediment. Litmus milk: Pinkish-brown ring; coagulated; peptonized, with alkaline reaction. Potato : Cream-colored growth, be- coming gray, wrinkled. Nitrites produced from nitrates. Soluble brown pigment formed. Sol- uble green pigment produced when freshly isolated. Starch is hydrolyzed. Aerobic . Optimum temperature 37°C. Source : Isolated once from upland California soil. Habitat: Soil. 4. Streptomyces californicus (Waks- man and Curtis) comb. nov. (Actinomyces californicus Waksman and Curtis, Soil Science, 1, 1916, 122.) Named for the State of California. Filaments with long, narrow, open spirals. Spherical to oval conidia from straight and spiral hyphae. Gelatin stab : Gray, moist, abundant surface growth. Liquefaction in 30 days. Synthetic agar: Spreading, vinaceous- colored growth. Aerial mycelium pow- dery, thin, light neutral gray. Starch agar: Growth spreading, pink center with colorless to gray margin. Glucose agar : Restricted, much folded, cream-colored growth, with sulfur-yellow tinge. Plain agar: Thin, restricted, yellowish to cream-colored growth. Glucose broth ; Solid cream-colored mass on surface, with pink tinge. Litmus milk: Faint, brownish surface growth; coagulated; peptonized in 40 days. Potato: Glossy, yellow to red growth, turning red-brown. Nitrites produced from nitrates. No soluble pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37°C. Source : Isolated once from California sandy loam. Habitat: Soil. 5. Streptomyces flaveolus (Waksman) comb. nov. (Actinomyces 168, Waksman, Soil Science, 8, 1919, 134; Actinomyces flaveolus Waksman, in Manual, 1st ed., 1923, 368; Actinomyces heimi Duchd, Actinomyces du groupe albus. Encyclo- pedic Mycologique, Paris, 6, 1934, 359.) From Latin flavus, yellow and eolus, diminutive ending; hence, somewhat yellow. FAMILY STREPTOMYCETACEAE 937 Xunierous closed and open spirals on all media. Conidia oval to elliptical. Gelatin stab: Liquefied; abundant, yellowish, spreading pellicle. Synthetic agar: Growth light sulfur- yellow turning to cadmium-j^ellow, pene- trating deep into medium. Aerial my- celium as white to ash-gray patches. Starch agar: White, spreading growth. Glucose agar: Restricted growth, sur- face folded, raised. Plain agar : White, glistening, wrinkled growth. Glucose broth: Thin, yellow pellicle. Litmus milk: Sulfur-yellow ring; co- agulated; peptonized, with faintly alka- line reaction. Potato: Abundant, wrinkled, cream- colored growth. Nitrites produced from nitrates. Soluble empire-yellow pigment formed. Starch is hydrolj'zed. Aerobic. Optimum temperature 25°C. Habitat: Soil. 6. Streptomyces bobiliae (Waksmanand Curtis) comb. nov. (Actinomyces bobili Waksman and Curtis, Soil Science, 1, 1916, 121.) From the name of a person. Mycelium with brandling hyphae. Few close spirals of a dextrorse type. Gelatin stab: Dense, cream-colored to brownish surface growth. Rapid lique- faction . Synthetic agar: Abundant, glossy, wrinkled, elevated, coral-red growth becoming deep red. Scant, white aerial mycelium. Starch agar: Restricted, tinely wrinkled, coral-red growth with hyaline margin . Plain agar: Restricted, glossy, gray growth, becoming brownish. Glucose broth : Round colonies in iluid. Flaky sediment. Litmus milk: Dark brown ring. Xo coagulation. Peptonized. Potato: Thin, yellowish growth, be- coming red, dry and wrinkled. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Source : Isolated once from adobe and garden soils. Habitat : Soil. 7. Streptomyces roseochromogenus (Jensen) comb. nov. {Actinomyces roseus Krainsky, Cent. f. Bakt., II Abt., 4/, 1914, 662; Waksman and Curtis, Soil Science, 1, 1916, 125; Waksman, Soil Science, 8, 1919, 148; not Actinomyces roseus Namyslowski, Cent. f. Bakt., I Abt., Orig., 62, 1912, 567; Nocardia rosea Chalmers and Christ opherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Actinomyces roseochromogenus Jensen, Proc. Linnean Soc. New So. Wales, 56, 1931, 359.) From Latin, producing rose color. Filaments with numerous open and closed spirals. Conidia 1.0 to 1.2 by 1.5 to 3.0 microns. Gelatin stab : Liquefaction, with small, cream-colored colonies in bottom of liquid. Synthetic agar: Thin, spreading, color- less growth. Aerial mj'celium thin, pale, brownish. Starch agar: Colorless, spreading growth. Glucose agar: Growth extensive, spreading, colorless, entire. Plain agar: White growth, becoming yellowish. Glucose broth : Cream-colored ring, with flaky sediment. Litmus milk: Brownish ring. No co- agulation. Peptonized in 10 to 15 days, becoming strongly alkaline. Potato: Much wrinkletl, brownish growth. Nitrites produced from nitrates. Purple pigment on egg media ; brown on gelatin. Starch is hydrolyzed. Aerobic. 938 MANUAL OF DETERMINATIVE BACTERIOLOGY Optimum temperature 37°C. Habitat : Soil . 8. Streptomyces griseolus (Waksman) comb. not'. (Actinomyces 96, Waksman, Soil Science, 8, 1919, 121 ; Actinomyces griseolus Waksman, in Manual, 1st eel., 1923, 369.) From Latin griseus, gray and eolus, diminutive ending; hence, some- what gray. Branching mycelium; no spirals ob- served. Conidia spherical or oval- shaped. Gelatin stab : Liquefied with yellowish, flaky pellicle and sediment. Synthetic agar : Colorless, thin, spread- ing growth, chiefly in the medium; sur- face growth limited almost entirely to the aerial mycelium. Aerial mycelium at first gray, later becoming pallid, neutral - gray. Starch agar: Grayish-brown growth, with dark ring. Glucose agar: Spreading growth, both on the surface and into the medium; center raised, cream-colored, turning dark. Plain agar: Brownish growth, with smooth surface. Glucose broth : Thick, brown ring. Litmus milk: Abundant growth, pink pellicle; coagulated; peptonized, becom- ing alkaline. Potato: Cream-colored growth, becom- ing black, spreading. Nitrites produced from nitrates. Faint brownish soluble pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Habitat: Soil. 9. Streptomyces erythreus (Waksman) comb. nov. (Actinomyces 161, Waksman, Soil Science, 8, 1919, 112; Actinomyces erythreus Waksman, in Manual, 1st ed., 1923, 370; Actinomyces krainskii Duchd, Encyclopedic Mycologique, Paris, G, 1934, 306.) From Greek erythrus, red. Mycelium fine, branching; numerous open spirals formed as side branches of the main hyphae. Gelatin stab : Abundant, dense, gray growth with pinkish tinge, chiefly on sur- face of liquefied medium. Synthetic agar: Spreading growth with irregular margin, developing deep into the medium; color at first white, later turning yellowish, agar around growth has a white, milky surface. Aerial myce- lium, thick, solid, white. Starch agar: Cream-colored, circular colonies, with faint greenish tinge. Glucose agar: Abundant, spreading, cream-colored growth, later turning brown chiefly on surface ; center raised, lobate margin. Plain agar: Cream-colored growth. Glucose broth: Abundant, cream- colored surface growth. Litmus milk : Yellowish surface zone ; coagulated; peptonized, becoming alka- line. Potato : Wrinkled, cream-colored growth, becoming yellowish. Nitrites produced from nitrates. Soluble purple pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Similar to Streptomyces erythrochromo- genes (Species No. 24) except that no brown soluble pigment is formed. Source : From California and Hawaiian soils. Habitat: Soil. 10. Streptomyces cellulosae (Krain- sky) comb. nov. {Actinomyces cellulosae Krainsky, Cent. f. Bakt., II Abt., U, 1914, 662.) From M. L. cellulosa, cellu- lose. Conidia almost spherical, 1.3 microns in diameter, often arranged in chains. Gelatin colonies: Circular, j'ellowish. Gelatin stab: Liquefied. Plain agar: White aerial mycelium. Ca-malate agar: Yellowish colonies; FAMILY STREPTOMYCETACEAE 939 gray aerial mycelium. Soluble j^ellow pigment formed. Glucose agar : Abundant growth, gray aerial mycelium. Soluble yellow pig- ment . Starch agar : Same as on glucose agar. Glucose broth: Coarse, flaky growth. Yellow pigment. Litmus milk: Peptonized. Potato : Light gray growth ; gray aerial mycelium. Nitrates show slight reduction. Strong diastatic action. Esculin is h}'- drolyzed. Cellulose is decomposed. Aerobic. Optimum temperature 30° to 35°C. Habitat: Soil. IL Streptomyces parvus (Krainsky) comb. nov. (Actinomyces parvus Krainsky, Cent. f. Bakt., II Abt., 41, 101-t, 662; Nocardia parva Chalmers and Chris- top herson, Ann. Trop. Med. and Para- sit., 10, 1916, 268.) From Latin parvus, small. Conidia more or less oval, 0.9 to 1.3 by 1.2 to 1.8 nucrons. Gelatin : Colonies yellow. Slow lique- faction. Ca-malate agar: Small, j-ellow colonies with light yellow aerial mycelium. Glucose agar : Same as on Ca-malate agar. Starch agar : Same as on Ca-malate agar. Glucose broth: Hemispherical colonies in bottom of tube. Litmus milk : Peptonized. Nitrate slightly reduced. Moderate diastatic action. Cellulose not decomposed. Aerobic. Optimum temperature. Source : Garden soil . Habitat: Soil. 12. Streptomyces malenconii (Duche) comb. nov. {Actinomyces malenconi Duch6, Encyclopedic Mycologique, Paris, 6, 1934, 353.) Named for Mr. :\Ialeugou from whom the original culture was obtained. Gelatin: Poor growth; liquefaction. Asparagine glucose agar : Rapid opaque growth, later becoming covered with white aerial mycelium; amber-colored pigment, dissolved in medium. Peptone agar : Cream-colored lobous growth, covered with whitish aerial my- celium. Asparagine glucose solution : Long, much branching filaments, 0.5 to 0.7 micron; somewhat heavier aerial myce- lium with a few irregular conidia; some flaky growth on bottom of tube; surface growth is cream-colored with rare white aerial mycelium; liquid becomes slightly j'ellow. Peptone solution : Whitish growth with yellowish soluble pigment. Milk : Surface growth with whitish aerial mycelium; slow peptonization, liquid becoming brownish-colored. Potato : Rapid growth with thin white mycelium; no soluble pigment. Coagulated serum : Radiating cream- colored growth covered with white aerial mycelium; slow liquefaction. No pigment on tyrosine medium. Source : Culture obtained from Mr. Malengon, an inspector in Morocco. 13. Streptomyces diastaticus (Krain- sky) comb. nov. {Actinomyces diastati- cus Krainsky, Cent. f. Bakt., II Bakt., 41, 1914, 662; Waksman and Curtis, Soil Science, /, 1916, 116.) From M. L. diastaticus, diastatic. Actinomyces roseodiastaticus Duche, Encyclopedic Mycologique, Paris, 6, 1934, 329 is said to differ from both Krain- sky's, and Waksman and Curtis' strains. Filaments may show fine, long, narrow spirals. Conidia oval, 1.0 to 1.2 by 1.1 to 1.5 microns. Gelatin stab : Liquefied with small, cream-colored flakes in liquid. Synthetic agar: Thin, gray, spreading 940 MANUAL OF DETERMINATIVE BACTERIOLOGY growth. Aerial mj'celium white, becom- ing drab gray. Starch agar: Thin, colorless, spreading growth. Aerial mycelium gray. Glucose agar : Yellowish, spreading growth. No aerial mycelium. Plain agar: Cream-colored growth. Thin aerial mycelium. Glucose broth: Gray ring with grayish colonies in bottom of tube. Litmus milk: Brownish ring; coagu- lated ; peptonized in 25 to 30 days, becom- ing faintly alkaline. Potato: Abundant, wrinkled, cream- colored growth with greenish tinge. Nitrites produced from nitrates. Brown to dark brown soluble pigment formed. Starch is hydrolyzed. Aerobic . Optimum temperature 37°C. Habitat: Soil. 14. Streptomyces fimicarius (Duche) comb. 710V. {Actinomyces fitiucanus Duche, Encyclopedie Mycologique, Paris, 6, 1934, 346.) From Latin fimus, tlung and caruSy loving. Gelatin : Punctiform colonies with whitish aerial mycelium; reddish soluble pigment. Liquefaction. Asparagine agar : Cream-colored growth with whitish aerial mycelium; reverse side, cream-colored to slight ochre. Czapek's agar : Yellowish masses of growth with yellowish-white aerial my- celium ; reverse side orange-colored ; faint yellowish soluble pigment. Peptone agar : Cream-colored growth with white aerial mj'celium; reverse side, yellowish. Asparagine solution : Vegetative fila- ments 0.5 to 0.6 micron long; branching aerial mycelium 0.8 to 1.0 micron, forming numerous conidia ; flaky growth produced on bottom; surface growth becomes cov- ered with a white aerial mycelium ; re- verse side, brownish-red. Czapek's solution: Cream-colored punctiform growth with yellowish aerial mycelium; no soluble pigment. Peptone solution : Whitish growth that flakes throughout liquid; yellowish pig- ment. Tyrosine medium: White growth with yellowish reverse ; yellowish soluble pig- ment. Milk : Colorless growth becoming cov- ered with whitish aerial mycelium; slow peptonization of milk which becomes rose- colored, finally changing to brownish-red. Potato : Cream-colored to yellowish growth with whitish aerial mycelium; reddish-brown pigmentation of plug. Coagulated serum : Cream-colored growth with whitish aerial mycelium; rapid liquefaction of serum. Distinctive characters : Abundant growth upon neutral and acid media; whitish aerial mycelium; marked odor; soluble brownish-red pigment. This spe- cies seems to form the transition type between the Actinomyces albus group and the Actinomyces chromogenus group. Habitat : Found abundantly in manure. 15. Streptomyces flavovirens (Waks- man) comb. nov. (Actinomyces 128, Waksman, Soil Science, 8, 1919, 117; Actinomyces flavovirens Waksman, in Manual, 1st ed., 1923, 352; Actinomyces albcviridis Duche, Encyclopedie ISIyco- logique, Paris, 6, 1934, 317.) From Latin flauus, yellow and virens, becoming green. Large masses of minute tufts ; the hy- phae coarse, straight, short, relatively unbranched, beaded; open spirals may be produced in certain substances. Conidia spherical, oval to rod-shaped, 0.75 to 1.0 by 1.0 to 1.5 microns. Gelatin stab: Yellowish-green surface pellicle, consisting of a mass of small colonies, on the liquefied medium. Synthetic agar : Growth spreading deep into the substratum, yellowish with greenish tinge. Aerial mycelium, gray, powdery . Starch agar : Greenish-yellow, spread- ing growth, developing deep into the medium. FAMILY STREPTOMYCETACEAE 941 Glucose agar; Restricted growth, de- veloping only to a very small extent into the medium, yellow, turning black, edge entire. Plain agar: Yellowish growth; the re- verse dark in center with yellowish zone and outer white zone. Glucose broth: Thick, sulfur-yellow pellicle or ring. Litmus milk : Cream-colored to brown- ish ring; coagulated; peptonized, becom- ing faintly alkaline. Potato : Sulfur-yellow, wrinkled growth. Only a trace of nitrite is formed from nitrates. Greenish-yellow soluble pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Habitat: Soil. 16. Streptomyces olivochromogenus (Bergey et al.) comb. nov. (Actinonujces chromogenus 205, Waksman, Soil Science, 8, 1919, 106; Actinomyces olivochromog- enus Bergey et al.. Manual, 2nd ed., 1925, 36S.) From Greek, producing an olive color. Filaments with numerous close spirals. Conidia oval or elliptical. Gelatin stab : Cream-colored, spreading surface growth. Rapid liquefaction. Synthetic agar: White, spreading growth. Aerial mycelium ash gray with brownish tinge. Starch agar: Transparent, spreading growth. Glucose agar: Abundant, natal brown to almost black growth, entire margin. Plain agar: Wrinkled, brown growth, becoming gray -green. Glucose broth: Thin, brown growth, flaky sediment. Litmus milk : Dark brown ring ; coagu- lated; peptonized, becoming alkaline. Potato : Small, wrinkled, black colonies. Faint traces of nitrites formed from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic . Optimum temperature 37 °C. Habitat: Soil. 17. Streptomyces diastatochromo- genes (Krainsky) comb. nov. {Actino- myces diastatochromogenes Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662.) From Greek, probably intended to mean producing both diastase and color. Conidia spherical or oval, about 1.2 microns. Gelatin colonies : Light gray-colored. Gelatin stab: Liquefied. Plain agar: Medium-sized colonies, with white to gray aerial mycelium. Ca-malate agar : Medium-sized colonies, colorless, with gray aerial mycelium. Glucose agar: Same as on Ca-malate agar. Starch agar : Same as on Ca-malate agar. Glucose broth : Flaky colonies in depth at first, later also over surface. Potato : Light gray colonies ; gray aerial nwcelium; medium colored black. Soluble brown pigment formed in gelatin. Weakly diastatic. Xo growth on cellulose. Tyrosinase formed. Aerobic. Optimum temperature 3o°C. Habitat: Soil. 18. Streptomyces flavochromogenes (Krainsky) comb. nov. {Actinomyces flavochromogenes Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662.) From Latin flavus, yellow and Greek, produc- ing color. Conidia oval, 1.7 microns. Gelatin colonies : Yellowish colonies. Gelatin stab : Slight liquefaction. Plain agar : Aerial mycelium formed late, at first white, later gray. Gray soluble pigment formed. Ca-malate agar : Colonies yellow with white aerial mycelium forming late. 942 MANUAL OF DETERMINATIVE BACTERIOLOGY Glucose agar: Brown soluble pigment formed. Starch agar: Yellow colonies, with white aerial mycelium. Glucose broth: Fine flakes, with small spherical colonies adherent to glass. Medium colored brown. Potato: Yellow colonies, with white aerial mycelium. Nitrites produced from nitrates. Weakly diastatic. Esculin acted upon. Slow growth on cellulose . Tyrosinase formed. Aerobic. Optimum temperature 35°C. Habitat: Soil. 19. Streptomyces antibioticus (Waks- man and Woodruff) comb. nov. {Actino- myces antibioticus Waksmau and Wood- ruff, Jour. Bact., 42, 1941, 232 and 246.) From Greek, antibiotic. Spore-bearing hyphae produced in the form of straight aerial hyphae. The conidiophores are arranged in clusters ; no spirals formed. The conidia are nearly spherical to somewhat elliptical. Gelatin : Dark brown growth on surface, with patches of gray aerial mycellium. Dark pigment produced, which gradually diffuses into the unliquefied part of the gelatin. Liquefaction at first very slow, later becoming rapid. Czapek's agar: Thin, whitish growth. Thin, gray aerial mycelium. Peptone media: Production of dark pigment at early stage of growth is very characteristic. Growth brownish, thin, with a yellowish-gray to yellowish-green aerial mycelium. Potato plug: Folded, brown-colored growth, with a thin black ring on plug, fading into a bluish tinge. No aerial mycelium. Carrot plug: Cream-colored to faint brownish growth. No aerial mycelium. No pigment. Litmus milk : Thick, brownish ring on surface of milk. Mouse-gray aerial my- celium with greenish tinge ; growth be- comes brown, especially in drier portions adhering to glass. No reaction change, no coagulation of milk, no clearing; whit- ish sediment at bottom of tube. Old cul- tures : Heavy growth ring on surface of milk, heavy precipitation on bottom; li- quid brownish to black in upper portion. Odor: Very characteristic soil odor. Antagonistic properties : Has a marked antagonistic effect on Gram-positive and Gram-negative bacteria, much more on the former than on the latter, as well as on actinomycetes. It is also active against fungi, which vary in degree of sensitivity. Produces a specific bacteriostatic and bactericidal substance known as actino- mycin (Waksman and Woodruff, Jour. Bact., 40, 1940, 581). Source : Isolated from soil on Escheri- chia coZt-washed-agar plate, using living cells of E. coll as the only source of avail- able nuti'ients. Habitat: Soil. 20. Streptomyces viridochromogenes (Krainsky) comb. nov. {Actinomyces vir- idochromogenes Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662; Waksman and Curtis, Soil Science, 1, 1916, 114.) From Latin, green and Greek, producing color. Filaments with numerous open spirals, 3 to 5 microns in diameter, occurring as side branches and terminal conidia, short ovals or spheres, 1.25 to 1.5 microns. Gelatin stab : Cream-colored surface growth, becoming greenish. Slow lique- faction. Synthetic agar : Spreading growth, cream-colored with dark center, becoming dark green; reverse yellowish to light cadmium. Aerial mycelium abundant, spreading, white, becoming light green. Starch agar: Circular, spreading, yel- lowish colonies. Glucose agar ; Abundant, spreading, wrinkled, gray growth, becoming black. Plain agar: Abundant, restricted, gray growth, with greenish tinge. FAMILY STKEPTOMYCETACEAE 943 Glucose broth: Dense, solid ring, brownish, becoming dark green. Litmus milk: Dark brown surface growth; coagulated; peptonized, with faintly alkaline reaction. Potato : Abundant, gray-brown growth. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Habitat : Soil . 21. Streptomyces purpeochromogenus (Waksman and Curtis) covih. nov. {Ac- tinomyces 'purpeochromogenus Waksman and Curtis, Soil Science, 1, 1916, 113.) From Latin, purple and Greek, producing color. Branching nn^celium and hyphae with few imperfect spirals. Conidia spheri- cal, 0.75 to 1.0 micron in diameter. Gelatin stab : Slow, brownish surface growth. Slow liquefaction. Synthetic agar: Slow, restricted, smooth, gray growth, becoming brown with purplish tinge; center raised. Margin yellow. Starch agar : Smal 1 , dark brown colonies . Glucose agar: Abundant, restricted, gray growth, becoming brown to dark brown. Plain agar: Gray to brownish growth, becoming dark brown, almost black. Glucose broth : Slight, flaky sediment. Litmus milk : Dark-brown ring ; coagu- lated; slowly peptonized, with faintly alkaline reaction. Potato: Restricted, orange to orange - red growth . Nitrites not produced from nitrates. Soluble dark brown pigment formed. Starch shows slight hydrolysis. Aerobic. Optimum temperature 25°C. Source : Isolated once from California adobe soil . Habitat : Soil. 22. Streptomyces phaeochromogenus (Conn) comb. nov. {Actinomyces pheo- chromogenus (sic) Conn, N. Y. State Agr. Exp. Sta. Tech. Bull. No. 60, 1917, 16.) From Greek, producing a brown color. Branching filaments and hyphae, spirals narrow, open, elongated, sinis- trorse . Gelatin stab : Abundant, spreading, cream-colored surface growth, becoming brown. Slow liquefaction. Synthetic agar: Colorless growth, be- coming brown to almost black. Aerial mycelium abundant, white with brownish shade. Starch agar: Spreading, brownish growth, becoming brown. Glucose agar : Restricted, much folded, brown growth . Plain agar : Thin, cream-colored growth, beconaing gray. Glucose broth : Dense, wrinkled pellicle. Litmus milk: Dark, almost black ring; coagulated, with slow peptonization, faintly alkaline reaction. Potato : Brown to almost black growth. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25''C. Source : Isolated from soil. Habitat : Soil. 23. Streptomyces aureus (Waksman and Curtis) comb. nov. {Actinomyces a7ire«.s Waksman and Curtis, Soil Science, 1, 1916, 124; not Actinomyces aurea Ford, Textb. of Bact., 1927, 220.) From Latin aureus, golden. Mycelium shows numerous spirals. Conidia spherical to oval, 0.6 to 1.0 by 0.8 to 1.4 microns. Gelatin stab : Fair, cream-colored sur- face growth, becoming brown, spreading. Liquefied. Synthetic agar : Thin, spreading, color- 944 MANUAL OF DETERMINATIVE BACTERIOLOGY less growth. Aerial mycelium thin, gray, powdery, becoming cinnamon drab. Starch agar : Thin, transparent, spread- ing growth. Glucose agar: Spreading, light orange growth, raised center, hyaline margin. Plain agar : Restricted, gray grov.th. Glucose broth: Thin, brownish ring; flaky sediment. Litmus milk: Black ring. No coagula- tion. Peptonization doubtful. Potato: Abundant, wrinkled, brown growth, becoming black. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Source : Isolated many times from a variety of soils. Habitat: Soil. 24. Streptomyces erythrochromogenes (Krainsky) comb, no v. (Actinomyces ery- throchromogenes Krainsky, Cent, f . Bakt., II Abt., Jtl, 1914, 662; Waksman and Cur- tis, Soil Science, /, 1916, 112.) From Greek, producing a red color. Conidia oval, about 2.0 microns long. Gelatin colonies: Slow growth. Gelatin stab: Liquefied. A soluble brown pigment formed. Plain agar: Brown soluble pigment. White aerial mycelium. Ca-malate agar : Colonies circular, with grayish-white margined aerial mycelium. Glucose agar: Red pigment formed. Starch agar : A soluble rose pigment on old cultures. Glucose broth: Abundant growth. Floating colonies, later a pellicle is formed. Brown soluble pigment. Potato: Gray aerial mycelium. Medium colored black. Nitrates show slight reduction. Weakly diastatic. No proteolytic enzyme formed. No growth in cellulose. Aerobic. Optimum temperature 30°C. Source : Soil and roots of Alnus (alder). Habitat: Soil. 25. Streptomyces lavendulae (Waks- man and Curtis) comb. nov. {Actino- myces lavendulae Waksman and Curtis, Soil Science, 1, 1916, 126.) FromM. L., lavender. Hyphae coarse, branching. Spirals close, 5 to 8 microns in diameter. Conidia oval, 1.0 to 1.2 by 1.6 to 2.0 microns. Gelatin stab : Creamy to brownish sur- face growth. Liquefied. Synthetic agar: Thin, spreading, color- less growth. Aerial mycelium cottony, white, becoming vinous-lavender. Starch agar : Restricted, glistening, transparent growth. Plain agar : Gray, wrinkled growth. Glucose broth: Abundant, flaky sedi- ment. Litmus milk: Cream-colored ring. No coagulation; peptonized, with strong al- kaline reaction. Potato: Thin, wrinkled, cream-colored to yellowish growth. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37°C. Certain strains of this organism pro- duce antibiotics. One such antibiotic, designated as streptothricin, is active both in vitro and in vivo against various Gram-positive and Gram-negative bac- teria, fungi and actinomycetes (Waks- man and Woodruff, Proc. Soc. Exp. Biol. I\Ied., 49, 1942, 207; Waksman, Jour. Bact., 46, 1943, 299). Source : Isolated once from orchard soil . Habitat: Soil. 26. Streptomyces reticuli (Waksman and Curtis) comb. nov. (Actinomyces reticuli Waksman and Curtis, Soil Sci- ence, 1, 1916, 118.) From Latin reticulum, a small net. Mycelium in whorls; spirals formed on FAMILY STREPTOMYCETACEAE 945 glucose agar are sinistrorse. Conidia spherical, 1.0 to 1.4 microns in diameter. Gelatin stab: Liquefied with small, brown flakes. Synthetic agar: Colorless growth, with yellowish tinge, becoming brownish, spreading. Aerial mycelium thin, white, cottony. Starch agar: Brownish-gray growth. Glucose agar: Restricted, brownish growth, center raised. Plain agar : Gra}^ wrinlded growth, be- coming brownish. Glucose broth : Sediment consisting of large colonies. Litmus milk: Reaction unchanged; co- agulated; peptonized. Potato: Gray growth, with black center. Nitrites produced from nitrates. Dark brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Source : From upland and adobe soils in California. Habitat: Soil. 27. Streptomyces rubrireticuli nom. nov. (Actinomyces reliculus -ruber Waks- man, Soil Science, 8, 1919, 146; Actino- myces reticulus Bergey et al., Manual, 2nd ed., 1925, 373.) From Latin ruber, red and reticulum, a small net. Branching filaments with both primary and secondary whorl formation. Spirals formed on glucose agar. Conidia oval- shaped. Gelatin stab : Surface growth yellow- ish-red to dragon-pink. Liquefied. Synthetic agar: Abundant, spreading growth, usually pink. Aerial mycelium thin, rose to pink. Starch agar : White growth with red tinge. Glucose agar: Abundant, spreading, rose-red, entire growth. Plain agar : Red growth, with yellowish margin, becoming red. Glucose broth : Thin, flaky sediment. Litmus milk : Abundant, red pellicle ; coagulated; peptonized. Reaction un- changed. Potato : Cream-colored growth, later pink to dark red. Nitrites produced from nitrates. Soluble dark brown pigment formed. Starch is hydrolyzed. Aerobic. Certain strains of this organism pro- duce an antibiotic. Source : Isolated from New Jersey orcliard and California upland soils. Optimum temperature 37°C. Habitat: Soil. 28. Streptomyces flavus (Krainsky) cornb. nov. (Actinomyces flavus Krain- sky, Cent. f. Bakt., II Abt., 41, 1914, 662; Waksman and Curtis, Soil Science, /, 1916, 118; not Actinomyces flavus San- felice. Cent. f. Bakt., I Abt., Orig., 36, 1905, 359.) From Latin ^ai^ws, yellow. Coarse filaments with branching hy- phae. Conidia formed by budding and breaking up of hyphae into oval forms. Gelatin stab: Small, yellowish masses on surface of liquefied medium. Synthetic agar : Circular, yellow or sul- fur-yellow colonies. Aerial mycelium straw-yellow. Starch agar: Spreading, cream-colored growth, with pink tinge. Glucose agar : Restricted, raised, folded, sulfur-yellow growth, center shading to brown. Plain agar: Gray, spreading, folded growth. Glucose broth : Small, white colonies in bottom of tube. Litmus milk: Coagulated; peptonized, becoming distinctly alkaline. Potato: Elevated, much wrinkled, greenish-olive growth. , Traces of nitrite formed. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. 946 MANUAL OF DETERMINATIVE BACTERIOLOGY Optimum temperature 25°C. Habitat: Soil. 29. Streptomyces ruber (Krainsky) comb. nov. (Actinomyces ruber Krain- sky, Cent, f . Bakt., II Abt., 41, 1914, 662 ; Waksman, Soil Science, 8, 1919, 149; not Actinomyces ruber Sanfelice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 355; No- cardia krainskii Chalmers and Chris - topherson, Ann. Trop. Med. and Para- sit., 10, 1916, 268.) From Latin ruber, red. Straight, branching mycelium, radiat- ing. A few spirals may be formed. Gelatin stab : Liquefaction, with yellow flakes. Synthetic agar: Abundant, spreading, red growth. Aerial mycelium abundant, cottony, chrome-orange. Starch agar : Abundant, spreading, red growth . Glucose agar: Restricted, abundant, entire, coral-red growth. Plain agar : Restricted, elevated, wrin- kled, olive-green growth. Glucose broth : Red ring, with spongy colonies on the surface. Litmus milk : Dark ring with red tinge ; coagulated; peptonized, with alkaline reaction . Potato: Elevated, wrinkled, greenish growth. Nitrites produced from nitrates. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Habitat: Soil. 30. Streptomyces citreus (Krainsky) comb. nov. {Actinoynyces citreus Krain- sky, Cent. f. Bakt., II Abt., 41, 1914, 662; Waksman and Curtis, Soil Science, 1, 1916, 99; not Actinomyces citreus Gas- perini,- Cent. f. Bakt., 15, 1894, 684.) FromM. L. citreus, lemon-yellow. Filaments with long, narrow open spirals. Conidia spherical to oval, 1.2 to 1 .5 by 1 .2 to 1 .8 microns. Gelatin stab : Yellowish, restricted sur- face growth. Liquefaction in 35 days. Synthetic agar: Abundant, spreading, raised, wrinkled, citron-yellow growth. Aerial mycelium covering surface ; citron- yellow. Starch agar : Abundant, yellowish- green growth. Glucose agar: Extensive, glossy, olive- yellow, entire growth; center elevated. Plain agar : Restricted, cream-colored growth . Glucose broth : Thin, wide, yellow ring ; flaky sediment. Litmus milk : Cream-colored surface growth; coagulated; peptonized, becom- ing alkaline. Potato: Yellowish growth, aerial my- celium white. Trace of nitrite production from ni- trate. The pigment formed is not soluble. Starch hydrolyzed. Aerobic. Optimum temperature 37°C. Habitat: Soil. 31. Streptomyces fulvissimus (Jensen) comb. nov. {Actinomyces fulvissimus Jensen, Soil Science, 30, 1930, 66.) From ha,tin fulvissimus, very yellow. Vegetative mycelium without any spe- cial characteristics; aerial mycelium of short, straight, often trifurcated hyphae, 1.0 to 1.2 microns broad; no spiral forma- tion; branches of hyphae break up into conidia, 1.0 to 1.2 by 1.2 to 1.5 microns. Gelatin : Vegetative mycelium narrow, smooth, yellowish-brown to red-brown; no aerial mycelium; no pigment; gelatin completely liquefied in 10 to 12 days. Nutrient agar : Good growth ; vegetative mycelium raised, finely wrinkled, deep red-brown ; no aerial mycelium ; brownish- yellow pigment. Czapek's agar : Good growth (one strain very scant), vegetative mycelium flat, narrow, first light golden, later deep orange to red-brown; aerial mycelium scant, sometimes almost absent, first FAMILY STREPTOMYCETACEAE 947 white, later light grayish-brown ; pigment very characteristic, bright golden to orange. Glycerol agar : Good growth ; vegeta- tive mycelium narrow, raised, smooth, golden to dark bronze; aerial mycelium scant, in patches, white to light cinna- mon-brown; pigment intensely golden to orange. Starch-casein agar : Good growth ; vege- tative mycelium spreading, folded, yel- lowish-brown ; aerial mj^celium abundant, smooth, lead -gray; pigment dull yellow to orange . Potato: Good growth; vegetative my- celium raised, much wrinkled, rust- brown; aerial mycelium absent or traces of white; pigment gray to faint lemon- yellow. Loeffler's blood serum : Vegetative mj^- celium red-brown; no aerial mycelium; yellowish pigment; no liquefaction. Distinctive characters : The character- istic golden pigment is formed in nearly all media in which the organism grows, but becomes most typical and attains its greatest brightness in synthetic agar media; it has indicator properties, turn- ing red in strongly acid solutions. The species is easilj^ recognized on agar plates by its bronze -colored colonies, sur- rounded b}^ haloes of bright j^ellow pig- ment. Source: Very common in Danish soils. Habitat: Soil. 32. Streptomyces gougeroti (Duche) cumb. nov. {Actinomyces gougeroti Du- che, Encyclopedic Mycologique, Paris, 6, 1934, 272.) Named for Prof. Gougerot, from whom the culture was obtained. Gelatin : Cream-colored colonies de- veloping slowly with faint aerial mj-- celium; no pigment; liquefaction. Plain agar : Cream-colored growth form- ing concentric ring with age, with brown- ish reverse; faint yellowish soluble pig- ment. Synthetic agar: Slow growth as puncti- form colonies ; cream-colored with smooth edge ; no aerial mycelium ; no soluble pig- ment. Peptone broth : Cream-colored ring on surface of medium with flakes throughout the medium; no soluble pigment. Synthetic solution : Submerged my- celium in the form of flakes, later forming a surface pellicle ; filaments of aerial my- celium 1 micron in diameter, with nu- merous conidia ; cream-colored growth ; no soluble pigment. Tyrosine medium : Good growth with white aerial mycelium; no soluble pig- ment. Litmus milk : Growth in the form of colonies which remain separated from one another ; also flakes in the bottom of the tube with bluish tinge on reverse of growth; milk turns blue in 10 to 12 days. Coagulated serum : Cream-colored growth covered with white aerial mj^ce- lium; rapid liquefaction of serum. Potato : Slow growth of a greenish tinge ; aerial mycelium; no black pigment. Distinctive character : Intermediate between Streptomyces albus with its abundant aerial mycelium and Actino- myces almquisti with its very scanty aerial mycelium. Source : Culture obtained from the col- lection of Prof. Gougerot. 33. Streptomyces violaceoniger (Waks- man and Curtis) comb. nov. {Actino- myces violaceus-niger Waksman and Curtis, Soil Science, 1. 1916, 111.) From Latin violaceus, violet and nigcr. black. Gelatin : Gray growth, with no produc- tion of aerial mj^celium. Gelatin around colonj^ rapidly liquefied, but without any change in color. Czapek's agar : Colony at first dark gray, turning almost black, 2 to 4 mm in diameter. Surface glossy, much folded with a very thin gray margin. A white to gray aerial mycelium is produced after the colony has well developed. A bluish- black pigment is produced at a later stage of its growth. The pigment slowly dis- solves in the medium, turning almost 948 MANUAL OF DETERMINATIVE BACTERIOLOGY black. Odor fairly strong. Microscopi- cally two types of mycelium are found : the thin, branching filaments of the sub- stratum, and the thick filaments of the aerial mycelium. The aerial mycelium fragments not very rapidly, producing a few conidia, spherical and oval, 1.2 to 1.5 by 1.2 to 2.3 microns. These often occur in chains. Czapek's solution: Colonies large, 2 to 3 mm in diameter, appearing at the bot- tom and surface of the solution, but none throughout the medium. Colonies blu- ish in color, with a regular margin. Me- dium not colored. Potato plug : Growth at first very slight, but after 48 hours develops into a yellow- ish-gray continuous thick smear which later turns brown, with a white aerial mycelium covering the growth. Medium not colored. Source : Isolated once from the upland California soil. Habitat: Soil. 34. Streptomyces griseus (Krainsky) comb. nov. {Actinomyces griseus Krain- sky, Cent. f. Bakt., II Abt., 41, 1914, 662. ) From M. L. griseus, gray. Branching filaments ; a few spirals have been observed. Conidia rod-shaped to short cylindrical, O.S bj' 0.8 to 1 .7 microns . Aerial mycelium greenish-gray. Gelatin stab : Greenish-yellow or cream- colored surface growth with brownish tinge. Rapid liquefaction. Synthetic agar : Thin, colorless, spread- ing growth, becoming olive buff. Aerial mycelium thick, powdery, water-green. Starch agar: Thin, spreading, trans- parent growth. Glucose agar: Growth elevated in cen- ter, radiate, cream-colored to orange, arose margin. Plain agar: Abundant, cream-colored, almost transparent growth. Glucose broth : Abundant, yellowish pellicle with greenish tinge, much folded. Litmus milk: Cream-colored ring; co- agulated with rapid peptonization, be- coming alkaline. Potato: Yellowish, wrinkled growth. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 37°C. Different strains of this organism pro- duce different antibiotics. One of these, streptomycin, was isolated in crystalline form. It is active against a large num- ber of bacteria and actinomycetes, but not against fungi and viruses. It is not very to.\ic to animals, and has found extensive application in the treatment of various diseases, mostly caused by Gram-negative bacteria and certain forms of tuberculosis. Source: Garden soil. ' Habitat: Soil. 35. Streptomyces griseofiavus (Krain- sky) comb. nov. {Actinomijces griseo- fiavus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662.) From M. L. griseus, gray and haiin flavus , yellow. Conidia oval, 1.2 microns. Gelatin colonies : Yellowish. Concentric rings. Gelatin stab : Rapidly liquefied. Plain agar: Colonies yellowish, with white aerial mycelium. Ca-malate agar : Large colonies covered with yellow to greenish-gray aerial my- celium. Glucose agar : White aerial mycelium is slowly formed. Starch agar : White aerial mycelium. Glucose broth : Flaky growth. Litmus milk: Peptonized. Potato: Yellowish growth, aerial my- celium gray. Nitrites produced from nitrates. Weakly diastatic. Acts on esculin. Grows well on cellulose. Aerobic. Optimum temperature 35°C. Habitat: Soil. FAMILY STREPTOMYCETACEAE 949 36. Streptomyces albidoflavus (Rossi Doria) comb. nov. (Streplotrix (sic) albido-flava Rossi Doria, Ann. d. 1st. d'Ig. sper. d. Univ. di Roma, 1, 1891, 407; Actinomyces albido-flavus Gas- perini, ibid., 2, 1892, 222; Streptothrix albido Chester, Man. Deterni. Bact., 1901, 365; Cladothrix albido-flava Mace, Traits Pratique de Bact., 4th ed., 1901, 1097; Nocardia albida Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271.) From Latin albidus, white and flavus, yellow. Description from Duch^, Encyclo- pedie Mycologique, Paris, 6, 1934, 294. Gelatin : Punctiform colonies with white aerial mycelium on surface of liquid; no soluble pigment; rapid lique- faction. Synthetic asparagine agar: Growth be- comes rapidly covered with white aerial mycelium, later becoming whitish- yellow ; brown on reverse side ; yellowish soluble pigment. Peptone agar : Cream-colored growth covered with fine white aerial mycelium; yellow soluble pigment. Tyrosine agar: Fine growth with orange-yellow on reverse side; medium becomes colored j^ellowish to yellowish- rose. Synthetic asparagine solution : Long branching filaments, 0.6 micron in di- ameter. Thicker aerial mycelium pro- ducing irregular spores ; flaky growth dropping to bottom of tube. Surface growth becomes covered with yellowish- white aerial mycelium; brownish on reverse side; soluble pigment yellowish. Peptone solution: Rapid, much folded growth, partly covered with white nw- celium on surface of medium; soluble yellow -ochre pigment. Milk : Rapid growth becoming covered with whitish aerial mycelium; never fully covering the surface ; no coagulation ; peptonization begins slowly and is com- pleted in 13 days, liquid becoming colored yellowish-orange . Coagulated serum : Cream-colored growth of surface becoming covered with white aerial mycelium ; rapid liquefaction of serum. Starch medium : Cream-colored growth rapidly colored with yellow aerial myce- lium; after 20 days growth becomes much folded ; greenish on reverse side ; slightly amber color in medium. This strain is closely related to Strepto- myces albus. Develops poorly on Czapek's medium without asparagine. Source: From dust. 37. Streptomyces poolensis (Tauben- haus) comb. nov. {Actinomyces poolen- sis Taubenhaus, Jour. Agr. Res., 13, 1918, 446.) Named for Prof. R. F. Poole, plant pathologist. Description from Waksman, Soil Sci., 8, 1919, 140. Fine, branching mycelium ; spirals usually not seen. Conidia oval to ellipti- cal. Gelatin stab: Liquefied, with small, brownish flakes in fluid. Synthetic agar : Thin, colorless, spread- ing growth. Aerial mycelium white to gray. Starch agar : Restricted, cream-colored growth. Glucose agar: Growth abundant, light brown, glossy, raised center, entire. Plain agar: Yellowish, translucent growth . Glucose broth : Thin, brownish ring. Litmus milk: Brownish ring; coagu- lated; peptonized, with strongly alkaline reaction. Potato : Thin, reddish-brown ; medium becoming purplish. Nitrites produced from nitrates. Faint trace of soluble brown pigment. Starch not hydrolyzed. Aerobic. Optimum temperature 37 °C. Source : Associated with disease of sweet potato. 950 MANUAL OF DETERMINATIVE BACTERIOLOGY 38. Streptomyces olivaceus (Waks- man) comb. nor. (Actinom3'ces 206, Waksman, Soil Science, 7, 1919, 117; Actinomrjces olivaceus Waksman, in Man- ual, 1st ed., 1923, 354.) From Latin, olive -colored. Small clumps, with straight and branching hyphae. No spirals on most media. Conidia spherical and oval, 0.9 to 1.1 by 0.9 to 2.0 microns. Gelatin stab : Liquefied with cream- colored, flaky, yellow sediment. Synthetic agar: Growth abundant, spreading, developing deep into medium, yellow to olive-ochre, reverse yellow to almost black. Aerial mycelium mouse - gray to light drab. Starch agar: Thin, yellowish-green, spreading growth. Glucose agar: Growth abundant, re- stricted, entire, center raised. Plain agar : White, glistening growth. Glucose broth : Sulfur-yellow ring. Litmus milk : Faint, pinkish growth ; coagulated; peptonized, becoming alka- line. Potato: Growth abundant, much wrinkled, elevated, gray, turning sulfur- yellow on edge. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Habitat: Soil. 39. Streptomyces lieskei (Duche) coynh. nov. {Aclinoviyces lieskei Duche, Encyclopedic M5^cologique, Paris, 6, 1934, 289.) Named for Prof. Lieske of Leipzig. Gelatin: Cream-colored growth be- coming covered with white aerial myce- lium; no soluble pigment. Rapid liquefaction. Plain agar : Cream-colored growth be- coming covered with white aerial myce- lium; yellowish soluble pigment. Synthetic agar : Cream-colored growth with delayed white aerial mycelium grow- ing from the edge toward the center; mycelium later yellowish. Reverse of growth yellowish to green. Dirty yellow to yellow-green soluble pigment. Synthetic solution : Long branching filaments 0.7 micron in diameter. Yel- lowish-white aerial mycelium does not readily produce spores ; flakes drop to the bottom of the tube. Peptone solution : Cream-colored colo- nies on surface with flakes in the liquid dropping to the bottom of the tube. Liquid becomes yellowish in color. Tyrosine medium : Rapid growth on surface with whitish-yellow aerial myce- lium; yellowish to orange-yellow soluble pigment. Milk: Cream-colored growth; colorless on reverse side; no aerial mycelium. Peptonization without coagulation. After 20 days the whole milk becomes a clear yellowish liquid. Coagulated serum : Clear-colored growth. Rapid liquefaction. Culture related to Streptomyces albo- jlavus and Streptomyces albidoflavus. 40. Streptomyces microflavus (Krain- sky) comb. nov. {Actinomyces micro- flavus Krainsky, Cent. f. Bakt.', II Abt., 4/, 1914, 662; Micromonospora microflava Duche, Encyclopedic Mycologique, Paris, 6, 1934, 29.) From Greek micnis, small, and Liat'in flavus, yellow. Conidia large, spherical to rod-shaped, often in pairs or chains, 2.0 by 2.0 to 5.0 microns. Gelatin colonies: Small, yellow. Gelatin stab: Liquefied. Plain agar: Yellow colonies, with rose- yellow aerial mycelium in 3 to 4 weeks. Ca-malate agar : Minute yellow colo- nies. No aerial mycelium. Glucose agar: A rose-yellow aerial my- celium develops in about 12 days. Starch agar : Same as on glucose agar. Glucose broth : Small spherical colonies in depth. Litmus milk: Peptonized. Potato: Yellow growth. No aerial mycelium. Nitrites produced from nitrates. FAMILY STREPTOMYCETACEAE 951 Strongly diastatic. Scant growth on cellulose . Starch is hj^drolyzed. Aerobic. Optimum temperature 25°C. Habitat : Soil. 41. Streptomyces cacaoi (Waksman) comb. 710V. (Actinomyces cacaoi Waks- man, in Bunting, Ann. Appl. Biol., 19, 1932, 515.) Named for the chocolate tree {Theohroma cacao). Long aerial mycelium with considerable spiral formation ; the spirals are long and open, not compact. Gelatin : Flocculent growth. Xo aerial mycelium. Rapid liquefaction. No pig- ment production. Nutrient agar: Brown-colored growth covered with tiny patches of ivory- colored aerial mycelium. Glucose agar: Thin yellowish growth, later turning reddish-brown; no soluble pigment; light gray to mouse-gray myce- lium, with white edge. Typical odor of streptomyces. Czapek's agar : Same as on glucose agar. Potato : Abundant brownish growth with white to mouse-gray aerial myce- lium. Biochemical characteristics : Strong proteolytic enzymes acting on casein and gelatin ; strong diastatic action, no sugar or dextrin left in 1 per cent starch solu- tion after a few days. Limited reduction of nitrate. Source : Three strains isolated from cacao beans in Nigeria. There were slight differences among the three strains ; the above description is of Strain L 42. Streptomyces novaecaesareae nom. 7tov. (Actinomyces violaceus-caeseri \Ya.ks- man and Curtis, Soil Science, 1, 1916, 111.) From Nova Caesarea, Latin name for the State of New Jersey. Filaments with both straight and spiral aerial hyphae ; spirals dextrorse. Conidia oval to elongate. Gelatin stab : Small, cream-colored sur- face colonies with slow liquefaction. Synthetic agar : Growth gray, becoming bluish, glossy, much wrinkled. Aerial mycelium appears late; white. Starch agar: Restricted, circular, blue- ish- violet colonies. Glucose agar : Restricted, gray growth, becoming red. Plain agar: Thin, cream-colored growth . Glucose broth: Fine, colorless, flakj- sediment . Litmus milk: Gray ring; coagulated; slow peptonization, becoming faintlj' alkaline. Potato : Growth cream-colored, wrinkled, turning yellowish. Nitrites produced from nitrates. Soluble purple pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Source : Isolated once from upland California soil. Habitat: Soil. 43. Streptomyces exfoliatus (Waks- man and Curtis) comb. nov. (Actino- myces exfoliatus Waksman and Curtis, Soil Science, 1, 1916, 116.) From Latin, stripped of foliage. Slightly wavy filaments with tendency to form spirals. Conidia oval, 1.0 to 1.5 by 1.2 to 1.8 microns. Gelatin stab : Cream-colored surface growth. Liquefied. Synthetic agar : Growth colorless, be- coming brown, smooth, glossy. Aerial mycelium in white patches over surface. Starch agar: Restricted, gray growth, becoming brown. Plain agar: Grows only in depth of medium. Glucose broth: Small, grayish colonies in depth. Litmus milk ; Cream-colored ring, soft coagulum in 12 days ; slow peptonization, becoming strongly alkaline. Potato : Growth somewhat wrinkled, gray, becoming brown. Nitrites produced from nitrates. Brown, soluble pigment formed. 952 MANUAL OF DETERMINATIVE BACTERIOLOGY Starch is hydrolyzed. Aerobic. Optimum temperature 37°C. Source : Isolated several times from adobe and upland soils in California. Habitat : Soil. 44. Streptomyces gelaticus (Waks- man) comb. nov. (Actinomyces 104, Waksman, Soil Science, 8, 1919, 165; Actinomyces gelaticus Waksman, in Manual, 1st ed., 1923, 356.) From M. L. gelaticus, gelatinous. Branching mycelium with open spirals. Gelatin stab : Liquefied with cream- colored flaky sediment. Synthetic agar: Growth colorless, spreading, chiefly deep into the medium. Aerial mycelium thin, white, turning grayish. Starch agar: Thin, spreading, cream- colored growth. Glucose agar: Abundant, spreading, white growth. Plain agar: Wrinkled, cream-colored growth only on the surface. Glucose broth : Thin, cream-colored pellicle; slight flaky sediment. Litmus milk : Pinkish ring ; coagulated ; peptonized with distinctly alkaline reac- tion. Potato: Growth abundant, much wrinkled, greenish, becoming black with yellowish margin. Nitrates show slight reduction to ni- trites. Soluble brown pigment formed. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Habitat: Soil. 45. Streptomyces rutgersensis (Waks- man and Curtis) comb. nov. (Actino- myces rutgersensis Waksman and Curtis, Soil Science, 1, 1916, 123.) Named for Rutgers University, New Brunswick, New Jersey. Branching filaments with abundant open and closed spirals ; hyphae fine, long, branching. Conidia spherical and oval, 1.0 to 1.2 microns, with tendency to bipolar staining. Gelatin stab : Cream-colored, spreading surface growth. Liquefied. Synthetic agar : Growth thin, colorless, spreading, becoming brownish to almost black. Aerial mj'celium thin, white, becoming dull -gray. Starch agar : Gray, spreading growth. Glucose agar : Abundant, brown myce- lium, becoming black with cream-colored margin. Plain agar: Thin, wrinkled, cream- colored growth. Litmus milk: Cream-colored ring; coagulated; slow peptonization, becoming alkaline. Potato: Abundant, white-gray, much folded growth. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Source : Isolated many times from a variety of soils. Habitat: Common in soil. 46. Streptomyces lipmanii (Waksman and Curtis) comb. nov. {Actinomyces lipmanii Waksman and Curtis, Soil Science, 1, 1916, 123.) Named for Prof. J. G. Lipman, New Jersey Agricultural Experiment Station. Straight, branching mycelium and hyphae. Conidia oval, 0.8 to 1.1 by 1.0 to 1.5 microns. Gelatin stab : Liquefied with cream- colored, flaky sediment. Synthetic agar : Growth abundant, raised, colorless, becoming light brown and wrinkled. Aerial mycelium white, turning gray. Starch agar: Transparent growth, be- coming dark with age. Glucose agar : Light yellow, irregular, spreading growth. Plain agar: Yellow, glossy, radiately wrinkled growth. Glucose broth: White ring, with abun- dant, colorless flaky sediment. FAMILY STREPTOMYCETACEAE 953 Litmus milk: Cream-colored ring; co- agulated; peptonization with alkaline re- action. Potato: Abundant, cream-colored, wrinkled growth. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Source : Isolated many times from a variety of soils. Habitat: Common in soil. 47. Streptomyces halstedii (Waksman and Curtis) comb. nov. (Actinonnjces halstedii Waksman and Curtis, Soil Science, 1, 1916, 124.) Named for a person. Branching mycelium and hyphae with close spirals. Conidia oval or rod- shaped, 1.0 to 1.2 by 1.2 to 1.8 microns. Gelatin stab: Liquefied, with small, cream-colored masses in bottom of tube. Synthetic agar: Growth abundant, heavy, spreading, raised, light, becoming dark, almost black. Aerial mycelium white, turning dull-gray. Starch agar : Abundant, brownish, glossy growth. Glucose agar : Growth spreading, color- less, wrinkled, center elevated, edge lichenoid, becoming brown. Plain agar: Restricted, wrinkled, cream-colored growth. Glucose broth : Small, colorless colonies in bottom of tube. Litmus milk: Cream-colored ring; co- agulated; peptonized, becoming alkaline. Potato : Growth abundant, moist, wrinkled, cream-colored with green tinge. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Optimum temperature 37 °C. Aerobic. Source : Isolated many times from the deeper soil layers. Habitat: Common in subsoil. 48. Streptomyces hygroscopicus (Jen- sen) comb. nov. (Actinomyces hygro- scopicus Jensen, Proc. Linn. Soc. New So. Wales, 56, 1931, 257.) From Greek, hygroscopic. Hyphae of vegetative mycelium 0.6 to 0.8 micron in diameter. Aerial liyphae long, tangled, branched, 0.8 to 1.0 micron in diameter; spirals numerous, sinis- trorse, narrow, usually short, only 1 or 2 turns, closed, typically situated as dense clusters on the main stems of the aerial hyphae. Conidia oval, 0.8 to 1.0 by 1.0 to 1.2 microns. Gelatin: Slow liquefaction. No pig- ment produced. Nutrient agar: Good growth. Vegeta- tive mycelium raised, wrinkled, glossy, cream-colored; later yellowish-gray with yellowish-brown reverse. Occasionally a scant white aerial mycelium. Sucrose agar : Good to abundant growth. Vegetative mycelium heavy, superficially spreading, folded, glossy surface, white to cream-colored, later sulfur-yellow to yellowish-graj^ with golden to light orange reverse. Soluble pigment of the same color. Aerial myce- lium scant, thin, white or absent. Glucose agar: Good growth. Vegeta- tive mycelium superficially spreading, surface granulated, cream-colored to straw-yellow, later dull chrome-yellow to brownish-orange. Aerial mycelium thin, smooth, dusty, white to pale yellowish- gray, after 1 or 2 weeks more or less abundantly interspersed with small, moist, dark violet-gray to brownish patches which gradually spread over the whole surface. Light yellow soluble pigment. Potato : Fair growth. Vegetative my- celium raised, wrinkled, cream-colored, later yellowish-gray to dull brownish. Aerial mycelium absent or trace of white. Milk: Completel}^ digested in 3 to 4 weeks at 30°C without any previous co- agulation. The reaction' becomes faintly acid (pH 6.0 or less) . Nitrates not reduced with sucrose as source of energy. Sucrose is inverted. 954 MANUAL OF DETERMINATIVE BACTERIOLOGY Starch is hydrolyzed. Cellulose is decomposed readily by some strains. Distinctive character: In this species, the aerial mycelium (which in other actinomycetes is strikingly hydrophobic) on certain media (glucose or glycerol asparagine agar) becomes moistened and exhibits dark, glistening patches. These patches, when touched with a needle, prove to be a moist, smeary mass of spores. This characteristic feature is not equally distinct in all strains. Source : Seven strains isolated from soils. Habitat : Soil . 49. Streptomyces fradiae (Waksman and Curtis) comb. nov. {Actinomyces f radii Waksman and Curtis, Soil Science, 1, 1916, 125.) From the name of a per- son. Straight, branching filaments and hy- phae. No spirals. Conidia rod- or oval- shaped, 0.5 by 0.7 to 1.25 microns. Gelatin stab : Cream-colored to brown- ish, dense growth on liquid medium. Synthetic agar: Smooth, spreading, colorless growth. Aerial mycelium thick, cottony mass covering surface, sea-shell pink. Starch agar : Spreading, colorless growth. Glucose agar: Growth restricted, glossy, buff-colored, lichenoid margin. Plain agar : Growth yellowish, becoming orange-yellow, restricted. Glucose broth: Dense, narrow, orange- colored ring; abundant, flak}?^, colorless sediment. Litmus milk : Faint, cream-colored ring; coagulated; peptonized, becoming alkaline. Potato: Restricted, orange -colored growth. Nitrites not produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 25°C. Source : Isolated once from adobe soil in California. Habitat: Soil. 50. Streptomyces alboflavus (Waks- man and Curtis) comb. nov. {Actinomyces albofiamts Waksman and Curtis, Soil Sci- ence, 1, 1916, 120.) From Latin albus, white and flavus, yellow. Straight, branching mycelium, with very little tendency to form spirals. Very few oval -shaped conidia formed. Gelatin stab : Abundant, colorless sur- face growth. Liquefaction occurs in 35 days. Synthetic agar: Growth glossy, color- less, spreading, becoming yellowish. Aerial mycelium white, powdery, with yellow tinge. Starch agar : Thin, yellowish, spread- ing growth. Glucose agar: Growth restricted, much-folded, creamy with sulfur-yellow surface . Plain agar : Restricted, cream-colored growth. Glucose broth : White, cylindrical colonies on surface, later flaky mass in bottom of tube. Litmus milk : Pinkish ring. No coagu- lation. Peptonized, becoming alkaline. Potato : Moist, cream -colored, wrinkled growth. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 37°C. Source : Isolated once from orchard soiL Habitat: Soil. 51. Streptomyces albosporeus (Krain- sky) comb. nov. {Actinomyces albo- sporeus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 649; Nocardia albosporea Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268; Waksman and Curtis, Soil Science, 1, 1916, 99.) From Latin albus, white and Greek spora, spore. FAMILY STREPTOMYCETACEAE 955 Straight, branching filaments with straight, branching hyphae, and occa- sional spirals. Conidia spherical or oval, 0.8 to 1.2 by 1.0 to 1.8 microns. Gelatin stab : Growth yellow, changing to red, with hyaline margin. Lique- faction in 35 days. Synthetic agar: Growth spreading, colorless with pink center, becoming brownish. Aerial mycelium white at first, later covering the surface. Starch agar: Growth thin, spreading, transparent, with red tinge. Glucose agar: Growth spreading, red, wrinkled, radiate, entire. Plain agar: Minute, cream-colored colonies . Glucose broth : Pinkish ring. Litmus milk: Scant, pink ring. No coagulation. No peptonization. Potato: Growth thin, spreading, wrin- kled, graj'', becoming brown with greenish tinge. Nitrites produced from nitrates. The pigment formed is not soluble. Starch is hydrolyzed. Aerobic. Optimum temperature 37 °C. Habitat: Soil. "52. Streptomyces flocculus (Duche) comb. nov. {Actinomyces flocculus Duche, Encyclopedie Mycologique, Paris, 6, 1934, 300.) From Latin, some- what woolly, referring to the appearance of the aerial mycelium. Gelatin: Very limited growth. Slow liquefaction. Asparagine glucose agar : Weak growth ; limited cream-colored colonies hardly raised above the surface of the medium ; occasionally abundant growth is produced with white aerial mycelium and colorless on reverse side. Czapek's agar: Cream-colored growth, later covered with white aerial mycelium ; no soluble pigment. Peptone agar: Cream-colored growth, later covered with white aerial mycelium ; no soluble pigment. Asparagine glucose solution : Branch- ing immersed filaments, 0.8 micron in diameter; aerial mycelium 1.0 by 1.2 microns with numerous conidia; flakes settle to the bottom of the tube. Peptone solution : Pointed colonies ; cream-colored on surface of medium. Tyrosine medium: Whitish growth without any pigment. Milk: Rose-colored growth; slow pep- tonization. Potato : Punctiform growth covered with white aerial mycelium ; faint yellow- ish pigment. Coagulated serum : Cream-colored growth; fine white aerial mycelium; slow liquefaction of serum. Source : Culture obtained from Mr. iVIalengon, an inspector in Morocco. 53. Streptomyces melanosporeus (Krainsky) comh. nov. {Actinomyces mel- anosporeus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662; Nocardia mclano- sporen Chalmers and Christopherson, Ann. Trop. I\Ied. and Parasit., 10, 1916, 268.) From Greek melas, black and spora, spore. Conidia almost spherical, 1.2 microns in diameter. Gelatin colony: Small, r,eddish colo- nies. Gelatin stab: Liquefied. Ca-malate agar: Colonies red, with black aerial mycelium. Glucose agar : Same as on Ca-malate agar. Starch agar : Same as on Ca-malate agar. Glucose broth: Flaky, orange-red colo- nies adherent to glass. Litmus milk: Peptonized. Potato : Red colonies with black aerial mycelium. Nitrites produced from nitrates. Weakly diastatic. Grows well on cellulose. Cellulose is decomposed. Aerobic. 956 MANUAL OF DETERMINATIVE BACTERIOLOGY Optimum temperature 25°C. Habitat: Soil. 54. Streptomyces melanocyclus (Mer- ker) comb. twv. (M icrocuccus melanocy- clus Merker, Cent. f. Bakt., II Abt., 31, 1911, 589; Acdtiomyces melanocyclus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662; Nucardia 7nelanocycla Chal- mers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268.) From Greek melas, black and cyclus, circle. Conidia almost spherical, 0.9 micron in diameter. Gelatin colonies : Growth poor. Gelatin stab : Rapid liquefaction. Ca-malate agar: Colonies small, flat, orange-red. Aerial mycelium black, oc- curring along the edges. Glucose broth: Same as on Ca-malate agar. Starch agar : Same as on Ca-malate agar. Glucose broth: Colorless, spherical colonies. Litmus milk: Peptonized. Nitrites produced from nitrates. Good diastatic action. Cellulose is decomposed. Aerobic . Optimum temperature 25°C. Habitat: Soil. 55. Streptomyces acidophilus (Jensen) comb. no/'. (Actinomyces acidophilus Jensen, Soil Sci., 25, 1928, 226.) From Greek, acid-loving. Vegetative mycelium profusely branched, hyphae 0.6 to 0.8 micron in diameter with homogeneous protoplasm and no visible septa. Aerial mycelium with hyphae 1.0 to 1.2 microns in di- ameter, somewhat branched, forming either very few or very numerous sin- istrorse spirals. Oval conidia 1.0 to 1.2 by 1.2 to 1.5 microns. Gelatin : After 10 days growth very scant, thin, colorless, semi-transparent. Slow liquefaction. Nutrient agar : No growth. Glucose agar: Good growth at 25°C. Substratum mycelium raised, somewhat wrinkled, colorless in young cultures. Aerial mycelium thin, white at first, later gray or yellowish-brown. Starch agar : Good growth at 25°C. Substratum mycelium flat, smooth, color- less. Aerial mycelium abundant, smooth, white. Czapek's agar : No growth. Plain broth: No growth. Milk: No growth. Potato: Growth good, raised, folded. No discoloration. Nitrites not produced from nitrates except a trace in two strains. Diastatic. Weakly proteolytic. Inversion of sucrose : Negative. Distinctive character : The ability to live in acid media only. Source : Four strains isolated from three acid humus soils. Habitat: Acid humus soils. 56. Streptomyces rubescens (Jarach) comb. nov. {Streptothri.r rubescens Javach, Boll. Sez. Ital. Soc. Intern. Microb., 3, 1931, 43.) From Latin rubescens, be- coming red. Gelatin: No liquefaction; limited non- pigmented growth. Glucose agar: Large number of small round colonies raised in the center and growing together, as well as deep into the medium; of a whitish opalescent color. Czapek's agar : Poor growth, becoming pigmented salmon-red, edge entire. Milk agar medium : Rose-coral-colored, thin growth with edge entire. Broth: Minute flakes, the liquid later becoming reddish-colored. Milk: No coagulation and no digestion; slight red coloration of milk. Potato plug: Reddish growth, not ex- tensive; opalescent surface. Source : From soil. 57. Streptomyces thermophilus (Gil- bert) comb. nov. {Actinomyces thermo- philus Gilbert, Ztschr. f. Hyg., 47, 1904, 383; not Actinomyces thermophilus Berestnew, Inaug. Diss., Moskow, 1897; FAMILY STREPTOMYCETACEAE 957 Nocardia thermophila Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271.) From Greek thermus, heat and phihis, loving. Description from Waksman, Umbreit and Gordon, Soil Sci., 47, 1939, 49. Hyphae straight, conidia formed. Gelatin: Liquefaction. Czapek's agar : At 28°C, deep colorless growth, thin white aerial mycelium; no soluble pigment. Starch agar : Yellowish growth with white-gray, powdery aerial mycelium. Milk: Proteolysis. Potato plug : Yellowish growth with no aerial mycelium, the plug usually being colored brown. Starch is hydrolyzed. No pigment produced on nutrient agar or gelatin. Temperature relations : Optimum 50°C. Good growth at 28°C. Usually no growth at 60°C. Some strains are in- capable of growing at 28°C, whereas others seem to grow well even at 65°C. Aerobic. Habitat: Soil, hay, composts. 58. Streptomyces thermofuscus (Waks- man, Umbreit and Gordon) comb. nor. (Actinomyces thermofuscus Waksman, Umbreit and Gordon, Soil Sci., 47, 1939, 49.) From Greek thermus, heat and Latin fuscus, dark. Presumably derived to mean heat -loving and dark in color. Hyphae spiral -shaped; conidia pro- duced. Gelatin: Liquefaction. At 50°C, a grayish ring is produced and soluble pig- ment is formed. At 28°C, growth with no soluble pigment. Czapek's agar : Poor growth at 28°C, deep-gray, with but little aerial myce- lium. At 50°C, growth dark to violet, with gray to lavender aerial mycelium and soluble brown pigment. Milk: Proteolysis. Potato : Abundant , dark-colored growth, no aerial mycelium, or few white patches, dark soluble pigment. Starch is hydrolyzed. Temperature relations : Good growth at 50° and 60°C. Will grow at 65°C. Faint growth at 28°C. Aerobic . Distinctive characters : This species is distinguished from Streptomyces thermo- philus by the brown-colored aerial myce- lium on synthetic media, spiral-shaped hyphae, and ability to grow readily at 65°C. Habitat: Soils and composts. 59. Streptomyces scabies (Thaxter) comb. nov. (Oospora scabies Thaxter, Ann. Rept. Conn. Agr. Exp. Sta., 1891, 153; Actinomyces scabies Gussow, Science, N. S. 39, 1914, 431.) From Latin scabies, scab. Wavy or slightly curved mycelium, with long branched aerial hyphae, show- ing a few spirals. Conidia more or less cylindrical, 0.8 to 1.0 by 1.2 to 1.5 mi- crons . Gelatin stab : Cream-colored surface growth, becoming brown. Slow lique- faction. Synthetic agar: Abundant, cream- colored, wrinkled, raised growth. Aerial mycelium white, scarce. Starch agar : Thin, transparent, spread- ing growth. Glucose agar: Restricted, folded, cream-colored, entire growth. Plain agar: Circular, entire colonies, smooth, becoming raised, lichenoid, wrinkled, white to straw-colored, opales- cent to opaque. Glucose broth: Ring in form of small colonies, settling to the bottom. Litmus milk : Brown ring with greenish tinge ; coagulated ; peptonized with alka- line reaction. Potato : Gray, opalescent growth, be- coming black, wrinkled. Nitrites produced from nitrates. Brown soluble pigment formed. Starch is hydrolyzed. Optimum temperature 37°C. Aerobic . The potato scab organism, like other 958 MANUAL OF DETERMINATIVE BACrERIOLOGY acid-fast organisms, can be selectively impregnated with carbol-auromin and when exposed to ultraviolet radiation fluoresces bright yellow. This technic confirms Lutman's conclusion that the hyphae are intercellular and grow within the middle lamellae (Richards, Stain Tech., 18, 1943, 91-94). Source : Isolated from potato scab le- sions. Habitat : Cause of potato scab ; found in soil. 60. Streptomyces ipomoea (Person and Martin) comb. nov. {Actinomyces ipo- moea Person and Martin, Phytopath., 30, 1940, 313.) From M. L. Ipomoea, a generic name. Conidia on glucose-casein agar : Oval to elliptical, 0.9 to 1.3 by 1.3 to 1.8 microns. Gelatin : After 25 days at 20°C, scanty growth, no aerial mycelium; no soluble pigment; liquefaction. Synthetic agar : Abundant growth, mostly on surface of medium, moderately wrinkled, olive -yellow. Nutrient agar : Moderate growth in the form of small, shiny, crinkled colonies both on the surface and imbedded in the medium, silver-colored. Starch agar : Growth moderate, smooth, deep in medium, ivory-colored. Aerial mycelium white with patches of bluish- green. No soluble pigment. Complete hydrolysis after 12 days. Milk : Growth in form of ring ; hydroly- sis, without visible coagulation. Potato : Growth moderate, light brown, shiny, wrinkled. No aerial mycelium. No soluble pigment. Nitrites are produced from nitrates. Starch is hydrolyzed. No growth on cellulose. Source : From diseased sweet-potato {Ipomoea sp.) tubers and small rootlets from several localities in Louisiana. 61. Streptomyces fordii (Erikson) comb. 710V. {Actinomyces fordii Erikson, Med. Res. Counc. Spec. Rept. Ser. 203, 1935, 15 and 36.) Presumably named for the surgeon who first secured the culture. Mycelium: Filaments of medium length, no spirals or markedly wavy branches. Short, straight, sparse aerial mycelium. Small oval conidia on potato agar and starch agar. Gelatin : No visible growth, slight softening in 20 days ; half-liquefied after 40 days. Agar: Small, creamy -golden, ring- shaped colonies, and heaped-up patches, becoming golden-brown in color and convoluted. Glycerol agar : Extensive, golden- brown, convoluted, thin layer. Serum agar : Golden-brown ring-shaped and coiled smooth colonies; no liquefac- tion. Ca-agar: Yellow, scale-like closely ad- herent colonies ; scattered white aerial mycelium. Blood agar : Innumerable small yellow- ish ring-shaped colonies; no hemolysis. Broth : Few flakes at first ; later abun- dant coherent puff ball growth. Synthetic sucrose solution : Moderate sediment of minute round white colonies. Synthetic glycerol solution : Light white fluffy colonies, minute and in clus- ters. Inspissated serum : Innumerable color- less pinpoint colonies ; scant white aerial mycelium; after 15 days colonies large, hollow on reverse side ; margin depressed ; no liquefaction. Dorset's egg medium: Minute, cream- colored, elevated colonies, becoming golden-brown, raised, convoluted. Milk: Coagulated; brownish surface ring. Litmus milk: No change in reaction. Potato plug : Yellowish growth in thin line, terminal portion tending to be piled up, scant white aerial mycelium at top of slant; after 12 days, growth abundant, golden-brown, confluent, partly honey- combed, partly piled up. Starch not hydrolyzed. Tyrosine agar: Reaction negative. Source : Human spleen in a case of acholuric jaundice. FAMILY STEEPTOMYCETACEAE 959 62. Streptomyces africanus (Pijper and Pullinger) cotnb. nov. {Nocardia africana Pijper and Pullinger, Jour. Trop. Med. and Hyg., 30, 1927, 153; Actinomyces ajricanus Nannizzi, in Pol- laci, Tratt. Micopat. Umana, 4, 1934, 8.) From Latin Africanus, relating to Africa. Description from Erikson, Med. Res. Counc. Spec. Rept. Ser. 203, 1935, IS. Unicellular branching mycelium form- ing small dense pink colonies with short straight sparse white aerial mycelium. Gelatin : Irregular pink flakes ; no liquefaction. Agar: A few flat pink discoid colonies. Glucose agar : Minute red discrete round colonies and piled up paler pink mass with thin white aerial mycelium. Glycerol agar: After 2 weeks, small heaped-up colorless masses with pink tinge around the colorless colonies, mar- gin depressed; after 3 weeks, abundant, piled up, pale pink growth. Ca-agar: After 1 week, small, round, colorless colonies with red centers, mar- gins submerged; after 2 weeks, growth bright cherry-red, confluent, with color- less margin. Dorset's egg medium : Small colorless blister colonies, partly confluent; be- coming wrinkled, depressed into medium ; slight liquefaction. Serum agar: Irregularly round, raised, wrinkled, colorless colonies; becoming dry, pink and flak}-; later piled up, brownish, friable. Inspissated serum : After one week, smooth, round, colorless colonies with submerged margin, in confluent patches pink and pitted into medium; after 2 weeks, medium broken up, slight lique- faction; after 3 weeks, liquid dried up, colonies umbilicated, raised, dry and friable. Broth: Small pink colonies embedded in coherent flocculent mass. Synthetic sucrose solution : Small pink granules in sediment after 1 week; colonies of medium size, coherent, after 3 weeks. Potato agar : Bright red growth, small round colonies with colorless submerged margins, and piled up patches with stiflE sparse white aerial mycelium. Litmus milk : Bright red surface growth, liquid unchanged after one month ; liquid opaque reddish-purple after 2 months; hydroh'zed, clear wine- red after 3 months. Source : From a case of mycetoma of a foot in South Africa. 63. Streptomyces gallicus (Erikson) comh. nov. {Actinomyces gallicus Erik- son, Med. Res. Counc. Spec. Rept. Ser. 203, 1935, 36.) From Latin gallicus, of the Gauls (French). Description from Erikson (loc. cit., p. 24). Mycelium shows lateral highly refrac- tive bodies which appear almost identical with the singly situated spores found in Micromonos-pora chalceae. Gelatin: Scant irregular pink growth; liquefaction very slow, only slight degree in 20 days. Agar : A few transparent minute pink colonies ; growth becomes partly con- fluent. Glucose agar : Xo growth. Glycerol agar : No growth. Czapek's agar: No growth. Coon's agar : Minute colorless to pink- ish colonies. Ca-agar: Glossy pink pinhead colonies. Potato agar : Pale pink, moist, granular growth. Serum agar: Pinpoint colonies, pink, shining. Blood agar : Abundant growth, minute, discrete, round, pink colonies, some aggregated in confluent narrow bands. No hemolysis. Dorset's egg medium: Minute colonies, becoming confluent, tangerine-colored. Inspissated serum : Abundant, pink, membranous growth, becoming reddish- brown; later discrete colonies at margin, clear on reverse side. No liquefaction. Broth : Pinkish flakes. Synthetic sucrose solution : A few fine white flocculi. 960 MANUAL OF DETERMINATIVE BACTERIOLOGY Synthetic glycerol solution : A few small round white colonies. Milk: Coagulated; peptonized; j'el- lowish-pink surface ring. Litmus milk: No coagulation or pep- tonization; no change in color. Potato plug: Very slow growth, a few minute translucent pink colonies after 16 days; after 21 days, considerable in- crease in number of colonies, still small and discrete. After 2 months, colonies 1 to 2 mm in diameter, bright coral, tending to be umbilicated and heaped up. T3'rosine agar: Reaction negative. Source : From blood culture in a case of Banti's disease. 64. Streptomyces pelletieri (Laveran) comb. nov. {Micrococcus pelletieri Lav- eran, Compt. rend. Soc. Biol., Paris, 61, 1906, 340; Oospora pelletieri Thiroux and Pelletier, Bull. Soc. path, exot., 5, 1912, 585; Nocardia pelletieri Thiroux, see Pinoy, Bull. Inst. Past., 11, 1913, 935; Discomyces pelletieri Brumpt, Precis de Parasitol., Paris, 2nd ed., 1913, 970; Actinomyces pelletieri Brumpt, ibid., 4th ed., 1927, 1204.) Named for M. Pelletier who first isolated this species. Description from Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 21. Thiroux and Pelletier (Bull. Soc. path, exot., 5, 1912, 585) considered that their cultures resembled Nocardia madurae, but they grew the organism only on Sabouraud's gelatin, on which it appeared in a constantly red, easily de- tachable form. Nocardia indica was regarded as identical by Pinoy, although in the original description by Laveran the organism was called Micrococcus pelle- tieri, owing to the fact that no mycelium was seen, merely coccoid bodies. No- cardia genesii Froes (Bull. Inst. Past., ^9, 1931, 1158) is described as closely allied, the distinction being founded upon the fact that the red grains were smaller in size and much more numerous, but no cultural details are given. Mycelium composed of slender straight and not very long filaments, forming small dense pink colonies with a few short straight isolated aerial branches. Gelatin : Slight liquefaction ; few pink flakes ; later almost completely liquefied. Agar : Minute colorless colonies and piled up pale pink masses. Glucose agar: Poor growth, a few minute pink colonies. Glycerol agar : Poor growth, a few moist pink colonies. Ca-agar : Colorless small colonies ; after 1 week, confluent skin, pink, buckled; medium discolored later. Coon's agar: Poor growth, cream- colored with pink center, mostly sub- merged. Potato agar : Colorless blister colonies ; after 3 weeks, colonies larger, showing concentric zones, submerged margins and occasional zone or tuft of white aerial mycelium, pinkish coloration. Dorset's egg medium: Abundant, wrinkled, pink skin with small discrete colonies at margin in six days; later surface rough, mealy; considerable lique- faction in 17 days. Serum agar: Moist cream-colored growth tending to be heaped up, discrete colonies at margin; becoming umbili- cated. Inspissated serum: Round, moist, colorless colonies. Blood agar : At first a few pinhead, cream-colored colonies, no hemolysis; later colonies dense, button-shaped, with narrow fringed margin. Broth: Small, minute, pink, clustered colonies. Synthetic sucrose solution: Small, pink colonies in sediment ; later minute col- onies adhering to side of tube. Milk: Soft curd; half -digested; pep- tonization complete in 20 days. Litmus milk : Pink surface growth, semi -solid, no color change; after 20 days, coagulum cleared, liquid purple. Potato plug: After one month growth sparse; yellowish-pink, irregularly piled up, portions with scant white aerial mycelium; after 6 months abundant highly piled up small rounded pink FAMILY STREPTOMYCETACEAE 961 masses, scant white aerial mycelium persistent. Source : From a case of crimson-grained mycetoma in Nigeria (E. C. Smith, Trans. R. Soc. Trop. Med. Hyg., 22, 1928, 157). Habitat : Human infections .so far as known . 65. Streptomyces listeri (Erikson) comb. 710V. {Actinomyces listeri Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for Dr. Joseph Lister, the father of antiseptic surgery. Description from Erikson (loc. cil., p. 23). Long slender filaments, many loosely wav}% forming a dense spreading my- celium which rapidly grows into a mem- brane on most media. Aerial mycelium very slow and inconstant in appearance, short and straight, conidia oval. Gelatin: Slight liquefaction; round white surface colonies; after 45 days, confluent skin, almost completely lique- fied. Agar: Smooth, round, moist, cream- colored, margin depressed, center ele- vated, closely adherent; becoming um- bilicated, with a myceloid margin. Glucose agar: Cream -colored, glisten- ing, pinpoint colonies ; later aggregated in convoluted skin. Glycerol agar: Abundant, moist, cream-colored growth, colonies elevated, piled up ; powdery white aerial mycelium. After 20 days, skin deeply buckled; colorless with exuded drops. Ca-agar: Poor growth, a slight biscuit- colored membrane. Potato agar : After one week, extensive growth, colorless submerged colonies, warted surface; dirty pink coloration after 2 weeks ; scant white aerial my- celium after 4 months. Dorset's egg medium: No growth. Blood agar: Small, round, cream- colored colonies, smooth translucent surface; no hemolysis. Serum agar: Small, irregular, moist, cream-colored colonies, tending to be heaped up; later somewhat transparent. Inspissated serum: Abundant growth, colorless shiny colonies, centrally ele- vated, becoming confluent. Broth: Small, round, white colonies in .sediment. Glucose broth: Small, white, nodular colonies; later abundant flocculi. Synthetic sucrose solution : Delicate white colonies in suspension and in sediment. Litmus milk: Coagulation. No change in reaction. Potato plug: Abundant, dull, brown- ish, wrinkled skin with white aerial mycelium; large, stellate, fluffy, white colonies in liquid at base. Source : From human material. Strain from Lister Collection. Habitat : From human infections so far as known. 66. Streptomyces upcottii (Erikson) comb. nov. (A new pathogenic form of Streptothrix, Gibson, Jour. Bact. and Path., 23, 1920, 357; Actinomyces upcottii Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for Dr. Harold Upcott, the surgeon who first secured the culture. Description from Erikson {loc. cit., p. 22). Filaments characteristically long, straight, much interwoven and ramified; typical unicellular mycelium, usually forming medium to large heavy carti- laginous colonies. Gibson states that the threads vary in thickness and show septa, but this has not been confirmed. A very slight transient aerial mycelium appeared on one agar slope, but this has not been repeated on any slide micro- culture on any medium. Slightly acid- fast. Gelatin : Abundant flocculent growth along streak, round cream-colored colonies on surface. Partly liquefied in 14 days ; complete liquefaction in 2 months . Agar: Smooth, shining, round, cream- colored colonies, margin submerged. 962 MANUAL OF DETERMINATIVE BACTERIOLOGY scant white aerial mycelium in one week ; colonies large (up to 10 mm in diameter), centers elevated, greenish tinge, very sparse aerial mycelium in two weeks ; the aerial mycelium disappears and large radial grooves appear in most colonies in 3 weeks. Glucose agar: Smooth, round, cream- colored colonies, margin depressed, cen- ters elevated, hollow on reverse side; later a coherent membranous growth, piled up, yellowish. Glycerol agar: Small, round, cream- colored, glistening colonies, heavy tex- ture, margins submerged; later, colonies umbilicated, tending to be piled up ; after 6 weeks, growth very much convoluted and raised, broad submerged margin, slightly reddish medium. Coon's agar: Small, radiating, white colonies, growth mostly submerged. Ca-agar: Small, colorless membranous growth with undulating margin; later, centrally depressed into medium. Potato agar: Poor growth, small, color- less blister colonies, medium slightly discolored. Dorset's egg medium: Round, flat, colorless, scale-like colonies, some marked by concentric rings and slightly hollowed in center; growth becomes yellow-brown. Serum agar : Large colonies (3 to 4 mm in diameter), colorless, granular, cen- trally elevated, depressed at margin, re- sembling limpets. Blood agar: Large drab heavily tex- tured colonies; no aerial mycelium; no hemolysis. Broth : Large coherent mass composed of fluffy colonies. Synthetic sucrose solution : Fair growth, minute white colonies. Carrot plug: Colorless, spreading, moist, wrinkled growth in six weeks; later a dull greenish-brown, moist, very much wrinkled and depressed skin. Source : From the spleen in a case of acholuric jaundice. Habitat : From human infections so far as known. G7. Streptomyces hortonensis (Erik- son) covib. nov. {Actinomyces horton Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for the Horton War Hospital at Epsom, England from which the culture was obtained. Description from Erikson {loc. cit., p. 22). Typical germination into very slow growing unicellular mycelium composed of long slender straight branching fila- ments. Very sparse straight aerial my- celium produced only once on potato. Non-acid -fast. Gelatin : Round cream-colored colonies on surface and a few mm below. No liquefaction. Agar : Very slow growth, a few smooth cream-colored coiled colonies in 19 days ; after 2 months, liberal, irregular, convo- luted growth. Glucose agar: Coiled and heaped up cream-colored translucent masses; after 2 months, growth rounded, elevated, ridged outwards from hollow center. Glycerol agar: Coiled, colorless, lus- trous patches, isolated colony with cen- tral depression. Serum agar : Poor growth, small amor- phous cream-colored mass. laspissated serum: Intricately coiled cream-colored growth. No liquefaction. Broth : Flakes . Synthetic sucrose solution : Poor growth, a few flakes. Synthetic glycerol solution: Delicate white flocculi at base. Litmus milk: Green surface growth, liquid hydrolyzed, partly clear purple; later decolorized, brown. Potato agar : Colorless blister colonies in one week ; dull green heaped and coiled mass after 3 weeks ; medium becomes slightly discolored. Potato plug: After 3 weeks, abundant, colorless, umbilicated, round colonies, some coiled in raised masses; later, liberal olive-green growth, piled up, dense, velvety gray-green aerial mycelium at top of slant, small round fluffy white colonies in liquid at base. FAMILY STREPTOMYCETACEAE 963 Source : From pus containing typical actinomycotic granules from parotid abscess. Habitat : From human infections so far as known. 68. Streptomyces gibsoaii (Erikson) comb. 710V. {Actinomyces gibsonii Erik- son, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for Prof. Gibson of Oxford. Description from Erikson (loc. cit., p. 15). Young growing mycelium branches profusely at short intervals ; later grows out into long frequently wavy filaments ; twisted hyphae also seen on water agar. Power of producing aerial mycelium apparently lost. Gelatin : Dull white flakes sinking as medium liquefies ; liquefaction complete in 12 days. Agar: Small, cream-colored, depressed, partly confluent colonies, becoming an extensive wrinkled cream-colored skin. Glucose agar: Cream-colored wrinkled membranous growth. Potato agar : Wrinkled glistening mem- branous growth. Serum agar : Small moist cream-colored colonies growing into medium. Dorset's egg medium: Small, round, smooth, colorless colonies with conically elevated centers. Inspissated serum: Innumerable color- less pinpoint colonies with scant white aerial mycelium at top; after 8 days, a coherent wrinkled skin with brownish- red discoloration at reverse, medium be- coming transparent; completely lique- fied, pigmented brown in 15 days. Blood agar: Yellowish confluent bands, irregularly wrinkled, with small discrete colonies, clear hemolytic zone. Broth : Sediment of fiocculi, some round and fan-shaped colonies. Synthetic sucrose solution: Very deli- cate white fiocculi. Potato plug : No growth. Starch not hydrolyzed. Milk: Coagulated; partly peptonized. Tyrosine agar: Negative reaction. Source : From the spleen in a case of acholuric jaundice. Injected into a mon- key, and reisolated. Habitat : From human infections so far as known. 69. Streptomyces beddardii (Erikson) comb. nov. {Actinoinijces beddardii Erik- son, Med. Res. Council Spec. Rept. Ser. •203, 1935, 36.) Presumably named for the surgeon who first secured the culture. Description from Erikson {loc. cit., p. 13). Rapidly growing, dense, spreading mycelium composed of very long slender filaments, many wavy or closely coiled, particularly on glucose agar; spirals less marked or lacking on poorer nutritive media like synthetic glycerol agar or water agar. Aerial mycelium sparse, short, straight on synthetic glycerol agar, much slower and more plentiful on glucose agar; later shows long, very fine spirals breaking up into small oval co- nidia; aerial h3'phae straighter and more branched with shorter conidiophores on starch agar. Non-acid-fast. Gelatin: Dull white flakes sinking to bottom as medium liquefies ; liquefaction complete in 8 days. Agar: Colorless, coherent, wrinkled, membranous growth with submerged margin; after 3 months, medium dis- colored, scant white aerial mycelium at top. Glucose agar: Wrinkled membranous growth; after 2 months, scant white aerial mycelium. Glycerol agar : Small, cream-colored, discrete colonies becoming confluent, under surface much buckled. Potato agar : Moist, cream-colored skin, convoluted, closely adherent. Ca-agar: Extensive, moist, cream- colored, wrinkled, membranous growth. Coon's agar: Scant, cream-colored, membranous growth. Starch agar : Spreading, colorless growth, considerable white aerial my- celium. 964 MANUAL OF DETERMINATIVE BACTERIOLOGY Blood agar: Hemolysis. Growth in uniformly striated colorless bands, occa- sional round colonies at margin. Dorset's egg medium: Extensive, very wrinkled, membranous growth, surface bright yellow. After 2 months, consider- able liquefaction. Serum agar: Wrinkled, glistening, cream-colored, membranous growth. Inspissated serum : Colorless smeary growth, reverse becoming transparent', starting to liquefy at base ; completely liquefied and brown in 12 days. Broth : Suspended and sedimented colorless flocculi, some small round colonies. Synthetic sucrose solution : Abundant white colonies in coherent mass near bottom of tube; large shell -shaped masses . Synthetic glycerol solution: At first, a few round white colonies in suspension; later, large branched feathery mass at bottom . Milk: Coagulated; later peptonized. Litmus milk : Medium deep blue, be- coming hydrolyzed to clear purple. Potato plug: Colorless moist membra- nous growth with scant white aerial my- celium at top of plug. Starch is hydrolyzed. Tyrosine agar: Reaction negative. Source : Human spleen in a case of spleenic anemia. Habitat : From human infections so far as known. 70. Streptomyces kimberi (Erikson) comb. nov. {Actinomyces kimberi Erik- son, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Presumably named for the surgeon who first secured the culture. Description from Erikson (loc. cit., p. 14). Mycelium of long straight profusely branching filaments forming circum- scribed colonies on all media with abun- dant production of short straight and branched aerial mycelium; small round conidia. Non-acid-fast. Gelatin: Liquefied. Smooth shining colonies becoming powdery white with aerial mycelium, floating on liquefied medium. No pigmentation. Agar : Smooth round moist cream- colored colonies, 1 mm in diameter; after 17 days, white powdery aerial mycelium. Glucose agar: Discrete cream-colored colonies becoming confluent, white aerial mycelium. Glycerol agar: M^ist cream-colored colonies becoming confluent, white aerial mj'^celium. Potato agar : Extensive growth covered by white powdery aerial mycelium; large colorless exuded droplets. Wort agar: Heavy brownish lichenoid colony; after 30 days, a white aerial mycelium. Ca-agar : Dull cream-colored scaly growth, covered by chalky white aerial mycelium. Coon's agar : Extensive growth, white aerial mycelium in annular arrangement. Czapek's agar: Small colonies covered with white aerial mycelium. Blood agar : Many large colonies, cream- colored, tough, smooth, glistening, with margin depressed; no hemolysis. Serum agar : Moist, cream-colored honeycombed skin, scant white aerial mycelium. Dorset's egg medium : Closely adherent scale-like colonies, centrally elevated, with white aerial mycelium. Lispissated serum : Rapid spreading growth, discrete round colonies at margin, completely covered with white aerial mycelium, colorless transpired 'drops ; slight softening at base. Broth : Small round colonies in sedi- ment in 2 days ; supernatant colonies with white aerial mycelium and large hollow flakes in sediment in 15 days; occasional reddish-brown coloration. Synthetic sucrose solution : Round white colonies at bottom; later small stellate colonies in suspension and a few supernatant with white aerial mycelium. Synthetic glycerol solution : Round white colonies at bottom; later coherent mulberry-like mass composed of fluffy FAMILY STREPTOMYCETACEAE 965 round portions; after 15 days, irregular wispy flocculi and large coherent mass. Milk: Coagulation; no peptonization; initial pinkish-brown ring descends until medium is dark brown throughout (2 months). Litmus milk : Blue coloration, hydro- Ij'zed to clear purple in 2 months. Starch not hydrolyzed. Tyrosine agar: Reaction negative. Source : Blood culture of a woman with acholuric jaundice. Habitat: From human infections so far as known. 71. Streptomyces somaliensis (Brumpt) comb. nov. (Indidla somaliensis Brumpt, Arch. Parasit., Paris, 10, 1906, 489; Discomyccs somaliensis Brumpt, Precis de Parasitologic, Paris, 2nd ed., 1913, 967; Indiellopsis somaliensis Brumpt, ibid.; Nocardia somaliensis Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 239; Strepiolhrix so7nalie7isis Miescher, Arch. Derm. Syph- ilis, 124^ 1917, 297; Actinomyces somalien- sis St. John-Brooks, Med. Res. Council Syst. of Bact., London, 8, 1931, 75.) Named for the country of origin, French Somaliland. Description from Erikson (Med. Res. Council Spec. Rept. Ser. 203, 1935, 17). Simple branching unicellular mycelium with long straight filaments, forming cir- cumscribed colony crowned with short straight aerial mycelium. Gelatin: Cream-colored colonies, me- dium pitted; complete liquefaction in 10 days ; hard black mass at bottom. Agar : Abundant yellowish granular growth wdth small discrete colonies at margin; later growth colorless, colonies umbilicated. Glucose agar: Poor growth, moist cream-colored elevated patch. Glycerol agar: Abundant growth, minute round to large convoluted and piled up masses, colorless to dark gray and black. Ca-agar : Round cream-colored colonies, depressed, umbilicated, piled up, thin white aerial mycelium; colonies become pale brown. Potato agar : Small round colorless colonies, zonate margin depressed, con- fluent portion dark greenish-black. Blood agar : Small dark brown colonies, round and umbilicated, piled up con- fluent bands, reverse red-black; hemol- ysis. Dorset's egg medium : Extensive color- less growth, partly discrete; becoming opaque, cream-colored, very wrinkled; later rough, yellow, mealy, portion liquid. Serum agar: Spreading yellow-brown skin, intricately convoluted. Inspissated serum : Cream-colored coiled colonies, medium pitted, trans- parent and slightly liquid. Broth : A few round white colonies at surface, numerous fluffy masses in sedi- ment ; later large irregular mass breaking into wisps. Synthetic sucrose solution: Minute round w^hite fluffy colonies in sediment ; after 17 days, scant wispy growth. Milk: Soft semi-liquid coagulum which undergoes digestion; heavy wrinkled surface pellicle, completely liquefied in 12 daj^s. Litmus milk : Soft coagulum, partly di- gested, blue surface ring; clear liquid in 12 days. Potato plug : Abundant growth, colonies round and oval, partly piled up in rosettes, frosted with whitish-gray aerial mycelium, plug discolored; after 16 days, aerial mycelium transient, growth nearly black. Although Streptomyces somalie7isis has been known for a long time, there has been until recently no detailed descrip- tions of the organism beyond the fact that it possesses a distinctly hard sheath around the grain which is insoluble in potash and eau de javelle. The rare oc- currence of septa and occasional inter- calary chlamydospores is reported by Brumpt (Arch. Parasit., 10, 1905, 562), but has not been confirmed by Erikson (loc. cit.). Chalmers and Christopherson (Ann. Trop. Med. Parasit., 10, 1916. 22-^) 966 MANUAL OF DETERMINATIVE BACTERIOLOGY merely mentioned the growth on potato as yellowish-white and lichenoid with- out describing any aerial mycelium. Balfour in 1911 reported a case but gave no data, and Fiilleborn limited his de- scription to the grain (Arch. Schiffs. Trop. Hyg., 15, 1911, 131). This species was first placed in Indiella, a genus of fungi, by Brumpt (1906, loc. cit.). Later Brumpt (1913, loc. cit.) proposed a new genus or subgenus, Indiellopsis , contain- ing the single species Indiellopsis somaliensis . Source : Yellow-grained mycetoma, Khartoum (Balfour, 4th Rept. Wellcome Trop. Res. Lab., A. Med., London, 1911, 365). Habitat : This condition has been ob- served by Baufford in French Somaliland, by Balfour {loc. cit.) in the Anglo-Egyp- tian Sudan, by Fiilleborn (loc. cit.) in German So. West Africa and by Chalmers and Christopherson {loc. cit.) in the Sudan. 72. Streptomyces panjae (Erikson) comb. nov. {Actinomyces panja Erikson, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for Dr. Panja who first secured the culture. Description from Erikson {loc. cit., p. 16). LTnicellular mycelium with slender branching filaments ; very small round colonies ; no aerial mycelium visible on any medium, but occasional isolated aerial branches. Non-acid-fast. Gelatin: Complete liquefaction in 4 days. Agar : Colorless irregularly piled up convoluted growth; after 1 month, easily detachable, brownish. Glucose agar : Small colorless coiled mass in 1 week; heaped up green growth in 2 weeks. Glycerol agar : Poor growth, scant colorless patch. Ca-agar : Colorless to pink spreading growth with minute discrete colonies at margin; after 2 weeks, bright red mass, buckled and shining, colorless sub- merged margin. Coon's agar : Small submerged colorless growth. Potato agar ; Small elevated convoluted colorless masses with purple tinge in center. Dorset's egg medium: Small round tough colorless colonies, margin well em- bedded; after 3 weeks, colonies elevated, warted, darkened, medium discolored and broken; slight degree of liquefaction, medium dark brown. Serum agar: Colorless, glistening, piled up, convoluted mass. Inspissated serum: Small round blister colonies and irregularly convoluted patches deeply sunk in pitted medium; after 2 weeks, medium transparent, slight degree of liquefaction. Broth : Flakes and minute colorless colonies. Glucose broth : Poor growth, scant flakes, pinkish. Synthetic sucrose solution : Pinkish flocculi; after 3 weeks, moderate growth, minute colorless colonies. Milk: Coagulation; pale green surface growth; mostly digested in 2 weeks. Litmus milk: Soft coagulum, color un- changed ; after 2 months, mostly digested, residue coagulum light purple. Source : From an ulcer of the abdominal wall, Calcutta. 73. Streptomyces willmorei (Erikson) comb. nov. {Actinomyces vnlhnorei Erik- son, Med. Res. Council Spec. Rept. Ser. 203, 1935, 36.) Named for Dr. Willmore who isolated the culture. Description from Erikson {loc. cit., p. 19). Germination usual, but growing uni- cellular mycelium frequently branches at very short intervals, presenting pe- culiar clubbed and budding forms with occasional separate round swollen cells which may represent the cystites of other writers. The filaments are character- isticallj' long, homogeneous, and much interwoven. Aerial mycelium is profuse FAMILY STREPTOMYCETACEAE 967 in most media, with a marked tendency to produce loose spirals (water and synthetic glycerol agar) with chains of ellipsoidal conidia. Thick aerial clus- ters may also be formed. Gelatin : Minute colorless colonies ; liquefaction. Agar : Heavy folded colorless lichenoid growth, rounded elevations covered with white aerial mycelium ; later, submerged margin, round confluent growth, aerial mycelium marked in concentric zones. Glucose agar : Colorless wrinkled con- fluent growth with smooth entire margin, large discrete colonies like flat rosettes ; after 4 months, scant white aerial my- celium. Glycerol agar : Round smooth cream- colored colonies, heavy texture, margin submerged, stiff sparse aerial spikes ; after 3 weeks, colonies large (up to 10 mm in diameter). Ca-agar : Spreading colorless growth, pitting medium, submerged undulating margin; verj^ scant white aerial mj'- celium. Coon's agar : Opaque white growth ex- tending irregularl}'^ (up to 3 mm) into medium, margin smooth and submerged, center raised, greenish tinge covered with white aerial mycelium; after 3 weeks, margin green, central mass covered by gray aerial mycelium. Potato agar: Fair growth, partly sub- merged, covered with grayish-white aerial mycelium; medium becomes dis- colored. Blood agar : Heavily textured small drab colonies, aerial mycelium microscop- ical ; no hemolysis . Dorset's egg medium: Large, round, colorless, scale-like colonies, radially wrinkled ; growth brownish, medium discolored in 2 weeks. Serum agar: Smooth colorless discoid colonies; marked umbilication after 2 weeks . Broth : Large fluffy white hemispherical colonies, loosely coherent. Synthetic sucrose solution : A few large round white colonies with smooth partly zonate margins, lightlj^ coherent in sedi- ment ; later smaller colonies in suspen.=!ion attached to side of tube. Milk: Coagulation; one-third pep- tonized. Carrot plug: Colorless raised colonies with powdery white aerial mycelium; after 1 month, very much piled up, aerial mycelium gra}'; after 2 months, super- abundant growth around back of plug, confluent, greatly buckled, all-over gray aerial mycelium. Source : Streptothricosis of liver (Will- more, Trans. Roj^ Soc. Trop. Med. Hyg., 17, 1924, 344). Habitat : From human infections so far as known. *Appendix: The following names have been used for species of Slreptoniyces. Many of them are regarded as new by their authors merely because they were isolated from a new tj^pe of lesion, or from some animal other than man. Others are inadequately" described species from air, soil or water. Relationships to other better described species are usually very obscure. Some of the species listed here may belong in the appendix to the genus Nocardia. Actinomyces aerugineus Wollenweber. (Arb. d. Forschungsinst. f. Kartoffelbau, 1920, 16.) From deep scab on potato. Actinomyces albidofuscns Neukirch. (Actinomyces albido fiiscus Berestnew, Inaug. Diss., Moskow, 1897; see Cent. f. Bakt., I Abt., 24, 1898, 707; Xeukirch, Inaug. Diss., Strassburg, 1902, 3.) From grain. Actinomyces albidus Duche. (Ency- clopedic Mycologique, Paris, 6, 1934, 266.) Actinomyces alhoatrus Waksman and * This appendix was originally prepared by Prof. S. A. Waksman and Prof. A. T. Henrici, May, 1943; it has been developed further by Mrs. Eleanore Heist Clise, Geneva, New York, August, 1945. 968 MANUAL OF DETERMINATIVE BACTERIOLOGY Curtis. (Soil Science, 1, 1916, 117.) From adobe soil . Actinomyces alhovlridis Duclid. (Eu- cyclopedie Mycologique, Paris, 6, 1934, 317.) Actinomyces albus (Rossi Doria) Gas ■ perini. (Streptotrix (sic) alba Rossi Doria, Ann. d. 1st. d'Ig. sper. d. Univ. di Roma, 1, 1891, 421 ; Streptothrix Nos. 2 and 3, Almquist, Ztschr. f. Hyg., 8, 1890, 189; Oospora doriae Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 251; Actinomyces bovis albv.s Gasperini, Atti Soc. Tosc. Scienz. Nat., P. V., 9, 1894; Gasperini, Cent. f. Bakt., 15, 1894, 685.) A general name applied to the most common streptomyces in air and water. Actinomyces albus asporogenes Berest- new. (Inaug. Diss., Moskow, 1897; see Cent. f. Bakt., I Abt., 24, 1898, 708.) Actinomyces albus var. ochraleucus WoUenweber. (Arb. d. Forschungsinst. fur Ivartoffelbau, 1920, 16.) Actinomyces albus - vulgaris Ciani. (Quoted from Baldacci, Boll. Sez. Ital. Soc. Internaz. di Microbiol., 9, 1937, 140.) Actinomyces almquisti Duche. (Duchi5, Encyclopedic Mycologique, Paris, 6, 1934, 278.) From culture labeled Actinomyces albus (Krainsky) Waksman and Curtis (Soil Sci., 1, 1916, 117; said to resemble Streptothrix No. 1, Almquist, Ztschr. f. Hyg., 8, 1890, 189). Actinomyces alni Peklo. (Cent. f. Bakt., II Abt., 27, 1910, 451.) From swellings of the roots of Alnus glutiyiosa. Actinomyces annulatus WoUenweber. (Arb. Forschungsinst. ftir Kartoffelbau, 1920, 10.) From dark-colored potato stem . Actinomyces (Streptothrix) amiulatus Beijerinck. (Folia Microbiologica, 1. 1912, 4.) Actinomyces aurea (du Bois Saint Sev^rin) Ford. (Streptothrix aurea du Bois Saint Sev^rin, Arch, de m^d. nav., 1895, 252; Nocardia aurea Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818; Oospora aurea Sartory, Champ. Paras. Homme et Anim., 1923, 818; Ford, Textb. of Bact., 1927, 220; not Actino- myces aureus Waksman and Curtis, Soil Science, 1, 1916, 124.) Possibly synony- mous witli Actinomyces aureus Lachner- Sandoval, Die Strahlenpilze, 1893, ac- cording to Lieske, Morphol. u. Biol, d. Strahlenpilze, Leipzig, 1921, 26. Found in conjunctivitis. Actinomyces bellisari Dodge. (Strep- tothrix alba Bellisari, Ann. Ig. Sperim., 14, 1904, 467; Oospora alba Sartory, Champ. Paras. Homme et Anim., 1923, 819; Dodge, Medical Mycology, St. Louis, 1935, 744.) Isolated in a ware- house in Naples from the dust of cereal coming from California. Actinomyces bovis var. nigerianus Erik- son. (Med. Res. Council Spec. Rept. Ser. 203, 1935, 20 and 36.) From strepto- thricosis of the skin of cattle in Nigeria. Actinomyces candidus (Petruschky) Bergey et al. (Streptothrix Candida (Gedanensis II) Petruschky, Verhandl. d. Kongr. f. innere Med., 1898; see Petruschky, in Kolle and Wassermann, Handb. d. path. Mikroorg., 2 Aufl., 5, 1913, 285 and 294; Nocardia Candida Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818; Discomyces candidus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 980; Bergey et al., Manual, 1st ed., 1923, 347.) From hu- man lung. Actinomyces carneus (Rossi Doria) Gasperini. (Streptotrix carnea Rossi Doria, Ann. 1st. d'Ig. sper. Univ. Roma, 1, 1891, 415; Gasperini, ibid., 2, 1892, 222; Oospora carnea Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 388; Cladothrix carnea Mace, Traits Pratique de Bact., 4th ed., 1901, 1096; Discomyces carneus Brumpt, Precis de Parasitol., 2nd ed., 1913, 976; Nocardia carnea Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818.) From air. Actinomyces carnosus Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces casei Bernstein and Mor- ton. (Jour. Bact., 27, 1934, 625.) Ther- mophilic. From pasteurized cheese. FAMILY STREPTOMYCETACEAE 969 Actinomyces cati (Rivolta) Gasperini. (Discomyces cati Rivolta, 1878; Gas- perini, Cent. f. Bakt., 15, 1894, 684.) Cause of a disease in a cat. Actinomyces cerebriformis Namyslow- ski. (Namyslowski, Cent. f. Bakt., I Abt., Orig., 62, 1912, 564; Streptothrix cerebriformis Chalmers and Christopher- son, Ann. Trop. Med. and Parasit., 10, 1916, 273; Nocardia cerebriformis Vuille- min, Encyclop^die Mycologique, Paris, B, 1931, 126.) From an infection of the cornea of the human eye. Actinomyces cereus. (Quoted from Lieske, Morphol. u. Biol. d. Strahlen- pilze, Leipzig, 1921, 33.) Actinomyces chromogenus (Gasperini) Gasperini. (Streptotrix nigra Rossi Doria, Ann. d. 1st. d'Ig. sper. d. Univ. di Roma, 1, 1891, 419; Streptotrix cromo- gcna (sic) Gasperini, according to Rossi Doria, zV/ew; Gasperini, ?6k/., 2, 1892, 222; Oospora chromogcncs Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 2, 1896, 389; Ctadothrix chromogencs Mace, Traite Pratique de Bact., 4th ed., 1901, 1075; Actinomyces niger Brumpt, Precis de Parasitol., Paris, 4th ed., 1927, 1206.) A general name for streptomyces from air producing a dark chromogenesis on protein media. Actinomyces cinereonigeraromaticus Neukirch. (Actinomyces cinereus niger aromaticus Berestnew, Inaug. Diss., Moskow, 1897; see Cent. f. Bakt., I Abt., 2^, 1898, 707; Neukirch, Inaug. Diss., Strassburg, 1902, 3, Nocardia cinereo- nigra Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271; Streptothrix cinereonigra aromatica, attributed to Berestnew by Chalmers and Christopherson, idem; Actinomyces cinereo-niger, quoted from Lieske, Mor- phol. u. Biol. d. Strahlenpilze, Leipzig, 1921, 33.) From grain. Actinomyces citreus Gasperini. (Gas- perini, Cent. f. Bakt., 15, 1894, 684; Streptothrix citrea Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 63; not Actinomyces citreus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662.) Actinoim/ces clavifer Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato and from soil. Actinomyces coroniformis Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From peat soil. Actinomyces craterifcr Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces cloacae Brussoff . (Cent, f . Bakt., II Abt., 49, 1919, 97.) From mud. Actinomyces cretaceus (Kriiger) Wol- lenvveber. (Oospcra cretacea Kriiger, Berichte der Versuchsstat. f. Zucker- rohrs, Kergok-Legal, 1890; Wollenweber, Arbeiten d. Forschungsinstitut fiir Kar- toffelbau, 1920, 16.) From potato scab. Actinomyces dicksonii Erikson. (Med. Res. Council Spec. Rept. Ser. 203, 1935, 17.) Actinomyces elaslica Sohngen and Fol. (Cent. f. Bakt., II Abt., 40, 1914, 92.) From garden earth. Actinomyces ferruginous Naumann. (Kungl. Svenska Vetenskapsakad. Hand!., I, 62, Part 4, 1921, 45.) From Aneboda region of Sweden. Deposits ferric hydroxide about the mycelial threads. Actinomyces filiformis (Boas) Nan- iiizzi. (Bacillus filiformis Boas, 1897; not Bacillus filiformis Tils, Ztschr. f. Hyg., 9, 1890, 294; not Bacillus filiformis Migula, Syst. d. Bakt., 2, 1900, 387; Nocardia filiformis Vuillemin, Encyclo- pedic Mycologique, Paris, 2, 1931, 132; Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 26.) From the human stomach. Actinomijces fimbriatus Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces flavogriseus T)uch6. (En- cyclopedie Mycologique, Paris, 6, 1934, 341.) From volcanic soils (Martinique). Actinomyces flavus Saufelice. (San- felice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 359; Streptothrix flava Sanfelice, ibid.; not Streptothrix flava Chester, Manual Determ. Bact., 1901, 362; not Actinomyces flavus Krainsky, Cent. f. 970 MANUAL OF DETERMINATIVE BACTERIOLOGY Bakt., il Abt., Al, 1914, 662.) From air. Actinomyces jlavus Millard and Burr. (Millard and Burr, Ann. Appl. Biol., IS, 1926, 601; not Actinonnjccs jlavus Sanfelice, Cent. f. Bakt., I Abt., Orig., S6, 1904, 359; not Actinomyces flaws Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662; not Actinomyces flavus Dodge, Medical Mycolog}^ St. Louis, 1G35, 752.) From scab on potato. Actinomyces Joersteri (Colin) Gas- perini. {Streptothrix Joersteri Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 3, 1875, 186 ; Cladothrix joersteri Winter, Die Pilzc, in Rabenhorst's Kryptogamen- Flora, I Abt., 1, 1884, GO; Kocardia Joersteri Trevisan, I generi e le specie dalle Batteriacee, 1889, 9; Oospora Joersteri Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 252; Gasperini, Cent. f. Bakt., 15, 1894, 684; Discomyces Joersteri Gedoelst, Les champignons parasites de I'homme et des animaux domestiques. Brussels, 1902, 176; Cohnistreptothrix Joersteri Pinoy, Bull. Inst. Pasteur, Paris, 11, 1913, 937.) The first strepto- myces to be described. Probably not identifiable. From an inflamed tear duct. Chalmers and Christopherson :Ann. Trop. Med. and Parasit., 10, 1916, 273) include Lcptolhrix oculorum Soro- kin, 1881 as a synonym of this species. Actinomyces Jusca Sohngen and Fol. (Cent. f. Bakt., II Abt., 40, 1914, 87.) From garden eartli. Actinoinyccs gabrilschewskii Xcukirch . (Actinomyces of Gabritschewsky, Berest- new, Inaug. Diss., Moskow, 1897; see Cent. f. Bakt., I Abt., 24, 1898, 708; Xeukirch, Inaug. Diss., Strassburg, 1902, 3.) From water. Actinomyces yedanensis (Lohlein) Ber- gey et al. (Strcptothrix yedanensis I, Scheele and Petruschky, Verhandl. d. Kongr. f. innere Med., 1897, 550; Strcpto- thrix riedanensis Lohlein, Ztschr. f. Ilyg., 6S, 1909, 11; Nocardia gedane7isis Chal- mers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 255; Disco- myces gedanensis Brumpt, Prdcis de Parasitol., Paris, 3rd ed., 1922, 984; Bergey et al., Manual, 1st ed., 1923, 347.) P^rom sputum of patient witli chronic lung disease. Actinomyces gibsoni Dodge. {Strepto- ihrix sp. Gibson, Jour. Path. Bact., 23, 1920, 357; Oospora sp. Sartory, Champ. Paras. Homme et Anim., 1923, 776; Dodge, Medical Mycology, St. Louis, 1935, 722.) See page 961. Actinomyces gracilis Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces graminearum Berestnew. (Berestnew, Inaug. Diss., Moskow, 1897; see Cent. f. Bakt.. I Abt., 24, 1898, 707; Nocardia graminarium (sic) Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 265; Strcptothrix graminarium Chalmers and Christopher- son, idem.) From grain. Actinomyces graminis Topley and Wilson. (Aktinomyces, Bostroem, Beitr. path. Anat. u. Path., 9, 1891, 1; Topley and Wilson, Princip. Bact. and Immun., 1st ed., 1, 1931, 250; Actino- myces hostroemi Baldacci, Boll. Sez. Ital. 8oc. Intermit. Microbiol., 9, 1937, 141.) From bovine actinomycosis. Actinomyces gruberi Terni. (Terni, Cent. f. Bakt., 16, 1894, 362; Nocardia gruberi Blanchard, in Bouchard, Traits Path. Gen., 2, 1896, 855; Strcptothrix grueberi (sic) Sanfelice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 356; Oospora gruberi, quoted from Nannizzi, Tratt. Micopat. Umana, 4, 1934, 51.) From soil. Produces several pigments on culture media. Actinomyces guignardi (Sauvageau and Radais) Ford. {Oospora guignardi Sau- vageau and Radais, Ann. Inst. Past., 6, 1892, 255; Ford, Textb. of Bact., 1927, 220.) From dust. Gasperini (loc. cit.) regards this as a possible synonym of Actinomyces chromogenus. Actinomyces halotrichis ZoBell and Upham. (Bull. Scripps Inst. Oceanog- raphy Univ. California, 5, 1944, 273.) From marine mud and kelp. Actinomyces heimi Duche. (Encyclo- FAMILY STREPTOMYCETACEAE 971 pddie JMycologique, Paris, 6, 1934, 359.) Actinomyces hoffmanni (Gruber) Gas- perini. {Micromijces hofmanni (sic) Gruber, Miinch. med. Wochnschr., 1891 ; also Arch. f. Hyg., 16, 1893, 35; Oospora hoffmanni Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 252; Gasperini, Cent. f. Bakt., 15, 1894, 684; Streptothrix hofmanni Kruse, in Fliigge, Die Mikro- organismen, 3 Aiifl., 2, 1896, 62; Clado- thrix hoffmanni Mace, Traits Pratique de Bact., 4th ed., 1901, 1081.) Patho- genic. See page 976. Actinomyces holmesi (Gedoelst) Xau- nizzi. {Discomyces holmesi Gedoelst, Champ. Paras. Homme et Anim., 1902; Xannizzi, Tratt. Micopat. Umana, 4, 1934, 49.) Actinomyces hominis Waksman. (Soil Science, 8, 1919, 129.) Culture received from K. IMeyer from Foulerton who iso- lated it in 1911 from an abscess of the palm. Waksman {loc. cit.) and Baldacci (Mycopathologia, 2, 1940, 160) regard this as identical with Bostroem's organ- ism (see Actinomyces graminis above) and Baldacci has renamed it Actinomyces innominatus. Actinomyces incanescens Wollenweber. (Arb. Forschungsinst. fiir Kartoffelbau, 1920, 16.) From the soil of potato fields near Berlin. Actinomyces intermedins (Kriiger) Wol- lenweber. (Oospora intermedia Kriiger, Berichte der Versuchsstat. f. Zucker- rohrs, Kergok-Legal, 1890; Wollenweber, Arb. d. Forschungsinst. flir Kartoffelbau, 1920, 16.) From the soil of potato fields near Berlin. Actinomyces interproximalis (Fennel) Ford. {Streptothrix interproximalis Fennel, Jour. Inf. Dis., 22, 1918, 567; Ford, Textb. of Bad.. 1927, 195.) From the mouth. Actinomyces invulnerabilis (Acostaand Grande Rossi) Lachner-Sandoval. (Cla- dothrix invulnerabilis Acosta and Grande Rossi, Cronica medicoquirurgica de la Habana, No. 3, 1893; see Cent. f. Bakt., 14, 1893, 14; Streptothrix invulnerabilis Kruse, in Fliigge, Die Mikroorganismen, 3 Aufl., 2, 1896, 64; Lachner-Sandoval, Ueber Strahlenpilze, Strassburg, 1898; Nocardia invulnerabilis Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271.) From river water. Actinomyces krausei (Chester) Ford. {Streptothrix aus Eiter, Krause, Miinch. med. Wchnschr., 46, 1899, 749 and Cent. f. Bakt., I Abt., 26, 1899, 209; also see Petruschky, in Kolle and Wassermann, 2 Aufl., 5, 1913, 267; Streptothrix krausei Chester, Manual Determ. Bact., 1901, 364; Nocardia krausei Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 263; Discomyces krausei Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 993; Ford, Textb. of Bact., 1927, 208.) From actinomy- cotic pus. Actinomyces lacerlae Terni. (Terni, L'Officiale Sanitario, 1896, 160; Strepto- thrix lacertae Foulerton, in Allbutt and Rolleston, Syst. of Med., 2, 1912, 309; Oospora lacertae Sartory, Champ. Paras. Homme et Anim., 1923, quoted from Xannizzi, Tratt. Micopat. Umana, 4, 1934, 51.) From grayish nodules in the liver of Italian lizards {Lacerta riridis and L. agilis). Actinomyces lathridii (Petruschky) Ford. {Streptothrix lathridii Pe- truschkj', Verhandl. d. Kong. f. innere Med., 1898; Ford, loc. cit., 205.) From the beetle, Lalhridius rugicollis. Actinomyces loidensis Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato Actinomyces luteo-roseus Sanfelice. {Actinomyces bovis luteoroseus Gasperini, Cent. f. Bakt., 15, 1894, 684; Sanfelice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 355.) Isolated from actinomycotic le- sion in cattle. Actinomyces marginatus Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces marinolimosus ZoBell and 972 MANUAL OF DETERMINATIVE BACTERIOLOGY Upham. (Bull. Scripps lust. Oceanog- raphy Univ. California, 5, 1944, 256.) From marine mud. Actinomyces melanoroseus Roisin. (Wisti Nauk Doslid. Kat. biol. Odessa, 1, 1929, 60.) ActinGrnyces metchnikoui (Sauvageau and Radais) Ford. {Oospura tnetchni- kowi Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 242; Ford, loc. cit., 220.) From water. Gasperini (loc. cit.) re- gards this organism as a possible synonym of Actinomyces chromogenus. Actinomyces muris ratti Lieske. {Streptoihrix ratti Schottmliller, Dermat. Wochnschr., 58, 1914, Supplement, 77; Streptoihrix muris-ratti Dick and Tunni- cliff, Jour. Inf. Dis.,:2.3, 1918, 186; Lieske, Morphol. u. Biol. d. Strahlenpilze, Leip- zig, 1921, 31; regarded as identical with Streptobacillus moniliformis Levaditi, Nicolau and Poincloux, Compt. rend. Acad. Sci. Paris, 180, 1925, 1188 by Topley and Wilson, Princip. of Bact.and Immun., 2nd ed., 1936, 274. The latter organism is regarded as identical with Haverhillia multiformis Parker and Hudson, Amer. Jour. Path., 2, 1926, 357 by Van Rooyen, Jour. Path, and Bact., 43, 1936, 469; Actinomyces muris Topley and Wilson, loc. cit.) From a case of rat-bite fever. Actinomyces musculorum Hertwig. {Actinomyces musculorum suis Duncker, Ztschr. f . Microskopie u. Fleischbeschau, 3, 1884. No. 3; Hertwig, Arch. f. wis- sensch. u. prakt. Thierheilk., 12, 1886, 365; Oospora musculorum suis Lehmann and Neumann, Bakt. Diag., 1 Aufl., 2, 1896, 383.) Seen in calcareous de- posits in the muscles of swine. Actinomyces myricac Peklo. (Cent. f. Bakt., II Abt., 27, 1910, 451.) From the roots of Myrica. Actinomyces from Neddeni, Namy- slowski. (Cent. f. Bakt., I Abt., Orig., 62, 1912, 564.) From the human eyelid. Actinomyces nigricaris Killiaii and Feh^r. (Ann. Inst. Past., 55, 1935, 620.) From desert soil. Actinomyces nigrijicans (Kriiger) Wol- lenweber. {Oospora nigrijicans Kriiger, Berichte der Versuchsstat. f. Zucker- rohrs, Kergok-Legal, 1890; Wollenweber, Arb. Forschungsinst. fiir Kartoffelbau, 1920, 16.) From potato scab. Actinomyces nitrogenes Sartory, Sar- tory, Meyer and Walter. (Bull. Acad. Med., Paris, 116, 1936, 186; also Ann. Inst. Past., 58, 1937, 684. ) From sputum. Actinomyces nivea. (Incorrectly at- tributed to Krainsky, 1914 by Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270.) Actinomyces nondiastaticus Bergey et al. (Var. b, Bergey, Jour. Bact., 4, 1919, 304; Bergey et al., Manual, 1st ed., 1923, 371.) From air. Actinomyces ochraceus Neukirch. (Ueber Actinomyceten, Strassburg, 1902, 4.) From soil. Actinomyces ochroleucus Neukrich. (Ueber Actinomyceten, Strassburg, 1902, 4.) From soil. Actinomyces odorifera Koelz. (Inaug. Diss., Kiel, 1934; Le Lait, 16, 1936, 154.) Actinomyces oligocarbophilus Lantzsch. (Lantzsch, Cent. f. Bakt., II Abt., 57, 1922, 309; Proactinomyces oligocarbophi- lus Krassilnikov, Bull. Acad. Sci., U. S. S. R., No. 1, 1938, 139.) Lantzsch regards this organism as identical with Bacillus oligocarbophilus Beijerinck and Van Delden, Cent. f. Bakt., II Abt., 10, 1903, 33 {Carboxydomonas oligocarbo- phila Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 311). Secures ' growth energy by oxidizing CO to CO2. From soil. See Manual, 5th ed., 1939, 81 for a description of the bacillary stage of this organism. Carboxyodomonas oligocarbo- phila Orla-Jensen is the type species of the genus Carboxydomonas Orla-Jensen (loc. cit.). Actinomyces orangico-nigcr. (Quoted from Lieske, Morphol. u. Biol. d. Strah- lenpilze, Leipzig, 1921, 33.) Actinomyces orangicus. (Quoted from Lieske, Morphol. u. Biol. d. Strahlen- pilze, Leipzig, 1921, 33.) Actinomyces pelogenes Sawjalow. FAMILY STREPTOMYCETACEAE 973 (Cent. f. Bakt., II Abt., 39, 1913, 440.) From mud containing hydrogen sulfide. Actinomyces pluricolor Terni. {Strep- tothrix pluricolor Fuchs; Terni, quoted from Gasperini, Cent. f. Bakt., 15, 1894, 684; Nocardia pluricolor Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268.) Actinomyces pluricolor diffundens Ber- estnew. (Inaug. Diss., Moskow, 1897; see Cent. f. Bakt., I Abt., 24, 1898, 708.) From air. Actinomyces praecox Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces praefecundus Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato and from soil. Actinomyces protea (Schiirmayer) Ford. {Oospora proteus and Streptothrix proteus Schiirmayer, Cent. f. Bakt., I Abt., 27, 1900, 58; Ford, loc. cit., 208.) From an abscess of the foot. Actinomyces pseudotuberculosae (Flex- ner) Brumpt. (Streptothrix pseudotuber- culosa Flexner, Jour. Exp. Med., 3, 1898, 438; Brumpt, Precis de Parasit., Paris, 4th ed., 1927, 1206.) Actinomyces pseudotuberculosus Leh- mann and Neumann. {Actinomyces atypica pseudotuberkulosa Haram and Keller, Cent. f. Bakt., I Abt., Ref., 42, 1909, 729; Lehmann and Neumann, Bakt. Diag., 5 Aufl., 2, 1912, 621; Nocardia pseudotuberculosis de Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 110.) Actinomyces purpureus Killian and Feher. (Ann. Inst. Past., 55, 1935, 620.) From desert soil. Actinomyces putorii (Dick and Tunni- cliff) Ford. (Streptothrix putorii Dick and TunniclifF, Jour. Inf. Dis., 23, 1918, 183; Ford, Textb. of Bact., 1927. 216.) From the 1)1 ood of a patient bitten by a weasel . Actinomyces pyogenes Lieske. (Eine neue Streptothrixspecies, Caminiti, Cent. f. Bakt., I Abt., Orig., U, 1907, 193; Streptothrix pyogenes Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Lieske, Morphol. u. Biol. d. Strahlenpilze, Leipzig, 1921, 32.) From air. Actinomyces radiatus Namyslowski. (Namyslowski, Cent. f. Bakt., I Abt., Orig., 62, 1912, 564; Streptothrix radiatus Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 273; Nocardia radiata Vuillemin, Encyclo- pedic Mycologique, Paris, 2, 1931, 126.) From an infection of the cornea of the human eye. Actinomyces rosaceus. (Quoted from Lieske, Morphol. u. Biol. d. Strahlen- pilze, Leipzig, 1921, 33.) Actinomyces roseodiastaticus Duch^. (Encyclopedie Mycologique, Paris, 6, 1934, 329.) Actinomyces roseus Namyslowski. (Ac- tinomyces sp. Lowensteins Klin. Mon- atsbl. f. Augenheilk., 48, 1910, 185; Namyslowski, Cent. f. Bakt., I Abt., Orig., 62, 1912, 567; not Actinomyces roseus Krainsky, Cent. f. Bakt., II Abt., 41, 1914, 662; Discomyces roseus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 981.) Actinomyces saharae Killian and Feh^r. (Ann. Inst. Past., 55, 1935, 621.) From desert soil. Actinomyces salmonicolor Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From sour soil. Actinomyces sampsonii Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces sanguinis Basu. (Ind. .Jour. Med. Res., 25, 1937, 325.) From the blood of a patient with bronchial pneumonia. Actinomyces sanninii Ciferri. (Quoted from Baldaeci, Boll. Sez. Ital. Soc. Internaz. di Microbiol., 9, 1937, 140.) Actinomyces setonii Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces spiralis Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From decaying grass. Actinomyces taraxeri cepapi (Schott- miiller) Ford. (Streptothrix taraxeri 974 MANUAL OF DETERMINATIVE BACTERIOLOGY cepapi Schottmiiller, Dermat. Wchnschr. , 68, 1914, Supplement, 77; Ford, loc. cit., 196.) From a case resembling rat-bite fever following the bite of a South African squirrel {Taraxerus cepapi). Actinomyces tenuis Millard and Burr. (Ann. Appl. Biol., 13, 1926, 601.) From scab on potato. Actinomyces thermodiastaticus Bergey et al. (Var. a, Bergey, Jour. Bact., 4, 1919, 301; Bergey et al.. Manual, 1st ed., 1923, 370.) From stomach contents of a rabbit. Actinomyces thermololcrans Stadler. (Arch. f. Hyg., 3S, 1899, 40.) From milk and butter. Actinomyces variabilis Cohn. (Cent. f. Bakt., I Abt., Orig., 70, 1913, 301.) From pus in the bladder' in a case of cystitis, and from the prostate. Actinomyces verrucosus Nadson. (Nadson, Die Mikroorganismen als geo- logische Faktoren I. Petersburg, 1903; quoted from Dorff, Die Eisenorganis- men, Pflanzenforschung, Jena, Hoft 16, 1934, 43; not Actinomyces verrucosus Adler, 1901, see Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934, 46.) From sea mud. Deposits ferric hydrox- ide about the mycelial threads. Actinomyces violaceus (Rossi Doria) Gasperini. {Streptotrix violacea Rossi Doria, Ann. d. 1st. d'Ig. sper. d. Univ. di Roma, 1, 1891, 411; Oospora violacea Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 252; Gasperini, Cent. f. Bakt., 15, 1894, 684; Cladothrix violacea Mac^, Traite Pratique de Bact., 4th ed., 1901, 1075; Nocardia violacea Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Discomyces viola- ceus Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 995.) From air and water. Actinomyces viridis (Lombardo-Pelle- grino) Sanfelice. (Streptothrix viridis Lombardo-Pellegrino, Riforma Med., 19, 1903, 1065; also see Cent. f. Bakt., I Abt., Ref ., 85, 1904, 761 ; Sanfelice, Cent, f. Bakt., I Abt., Orig., 86, 1904, 355.) From soil. Actinomyces viridis Millard and Burr. (Millard and Burr, Ann. Appl. Biol., .ZS, 1926, 601; not Actinomyces viridis San- felice, Cent. f. Bakt., I Abt., Orig., 86, 1904, 355; not Actinomyces viridis Duch^, Encyclopedic Mycologique, Paris, 6, 1934, 311.) From scab on potato. Actinomyces xanthostromus Wollen- weber. (Arb. Forschungsinst. f. Kar- toffelbau, 1920, 16.) Actinomyces wedmorensis Millard and Burr. (Ann. Appl. Biol., 18, 1926, 601.) From peat soil. Asteroides lieskeyi Puntoni and Leo- nardo (Boll, e Atti d. R. Accad. Med. di Roma, 61, 1935, 94.) A renaming of Actinomyces lieskey, a culture whose source was unknown. This may pos- sibly be the same as Actinomyces lieskei Duche (see Streptomyces lieskei). Cladothrix odorifera Rullmann. (Rull- mann, Inaug. Diss., Munich, 1895; see Cent. f. Bakt., I Abt., 17, 1895, 884 and Cent. f. Bakt., II Abt., 2, 1896, 116; Oospora odorifera Lehmann and Neu- mann, Bakt. Diag., 1 Aufl., 3, 1896, 392; Actinomyces odorifer Lachner-Sandoval, Ueber Strahlenpilze, 1898, 65; Strepto- thrix odorifera Foulerton and Jones, Trans. Path. Soc. London, 58, 1902, 112; Nocardia odorifera Castellani and Chal- mers, Man. Trop. Med., 2nd ed., 1913, 818.) From sputum in a case of chronic bronchitis. Cladothrix placoides Kliglcr. (Lcpto- Ihrix placoides alba Dobrzyniecki, Cent, f. Bakt., I Abt., 21, 1897, 225; Kligler, Jour. Allied Dental Soc, 10, 1915, 141, 282 and 445; Leptotnchia placoides Bergey et al., Manual, 3rd ed., 1930, 458.) From a tooth canal. For a description of this species see Manual, 5th ed., 1939, 829. The description indicates that this organism belongs to Nocardia. or Streptomyces. C occobacillus psevdo - actinoynycosis polymorphus Berestneff. (Berestneff, 1898, quoted from Chalmers and Chris- topherson, Ann. Trop. Med. and Para- sit., .?0, 1916, 273.) Cohnistreptothrix americana Chalmers and Christopherson. {Streptothrix sp. FAMILY STREPTOMYCETACEAE 975 Bloomfield and Bay ne- Jones, Johns Hopkins Hosp. Bull. 26, 1915, 230; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 273; Actinomyces americanus Dodge, Medical Mycolog3^ St. Louis, 1935, 716.) From a liver abscess. Cohnistreplothrix viisri Carpano. (Riv. di Parasit., No. 2, 1937, 107.) From human dermatosis in Egypt. Cohnistreptothrix silberschmidtn Chal- mers and Christopherson. (Streptothrix, Silberschmidt, Cent. f. Bakt., I Abt., £7, 1900, 486; Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 273; Nocardia silberschmidii Froilano de Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, HI; Actinomyces silherschmidti Dodge, INledical Mycology, St. Louis, 1935, 711.) From cases of dacryocystitis. Discomyces decussalus Langeron and Chevallier. (Langeron and Chevallier, Compt. rend. Soc. Biol., 72, 1912, 1030; Nocardia decussata Castellani and C^hal- mers, Man. Trop. Med., 2nd ed., 1913, 817; Oospora decussata Sartory, Champ. Paras. Homme et Anim., 1923, 825; Actinomyces decussalus Brumpt, Precis de Parasitol., 4th ed., 1927, 1206.) From dry, scaly lesions. Not considered path- ogenic. Nocardia chalmersi de Mello and Fer- nandes. (De Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 130; Actinomyces chalmersi Dodge, Medical Mycology, St. Louis, 1935, 734.) From the saliva of a horse. Nocardia Christopher soni de Mello and Fernandes. (De Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 130; Actinomyces christophersoni Dodge, Med- ical Mycology, St. Louis, 1935, 723.) From the air. Nocardia citrea Chalmers and Chris- topherson. (Ann. Trop. Med. and Para- sit., 10, 1916, 270.) A blanket name proposed to include Actinomyces griseo- flavus Krainsky, Actinomyces flavus Kra- insky, Streptothrix flava Sanfelice and Streptothrix flava Bruns. Nocardia cruoris Macfie and Ingram. (Macfie and Ingram, Ann. Trop. Med. and Parasit., 15, 1921, 283; Discomyces cruoris Brumpt, Precis de Parasitol., Paris, 3rd ed., 1922, 984; Oospora cruoris Sartorj', Champ. Paras. Homme et Anim., 1923, 809; Actinomyces cruoris Brumpt, ibid., 4th ed., 1927, 1195.) From blood. Nocardia dichotoma (Mace) Chalmers and Christopherson. (Cladothrix dicho- toma Mace, Compt. rend. Acad. Sci. Paris, 6, 1888, 1622; not Cladothrix dicho- toma Cohn, Beitr. z. Biol. d. Pflanzen 1, Heft 3, 1875, 185; Chalmers and Chris- topherson, Ann. Trop. Med. and Para- sit., 10, 1916, 270.) Nocardia ferrvginea Trevisan. (Bakterium bei Chorea St. Viti, Naunyn, Mittheil. aus der Med. Klinik zu Konigs- berg, 1888, 292; Trevisan, I generi e le specie delle Batteriacee, 1889, 9; Actino- myces Jerrugineus Gasperini, Cent. f. Bakt., 15, 1894, 684.) From pia mater in a ca.se of St. Vitis' dance. Nocardia gartcni (Brumpt) Castellani and Chalmers. {Cladothrix liquefaciens No. 2, Garten, Deutsche Ztschr. f. Chirurg., U, 1895, 432; Discomyces gartcni Brumpt, Precis de Parasitol., Paris, 1st ed., 1910, 860; Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818; Oospora garteni Sartorj'-, Champ. Paras. Homme et Anim., 1923, 778; Actinomyces garteni Brumpt, loc. cil., 4th ed., 1927, 1191; Actinomyces liquefaciens Ford, Textb. of Bact., 1927, 202.) From cases of human actinomy- cosis. Nocardia goensis de Mello and Fer- nandes. (De Mello and Fernandes, Mem. Asiatic Soc. Bengal, 7, 1919, 130; Actinomyces goensis Dodge, Medical Mycology, St. Louis, 1935, 723.) From lesions of vitiligo. Saprophytic. Nocardia liguire Urizer. (Urizer, 1904; Actinomyces liguire Nannizzi, in PoUacci, Tratt. Micopat. Umana, 4, 1934, 49.) Nocardia liquefaciens (Hesse) Castel- 976 MANUAL OF DETERMINATIVE BACTERIOLOGY lani and Chalmers. {Cladothrix lique- Jaciens Hesse, Deutsche Ztschr. f. Chirurg., ^1, 1895, 432; Discomyces liqiie- faciens Brumpt, Precis de Parasit., Paris, 1st ed., 1910, 860; Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 818; Streptothrix liquefaciens Chal- mers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 233, 265; Oospora liquefaciens Sartory, Champ. Paras. Homme et Anim., 1923, 778; Actinomyces liquefaciens Brumpt, loc. cit., 4th ed., 1927, 1192.) From an in- guinal abscess. Nocardia microparva Chalmers and Christopherson. (Ann. Trop. Med. and Parasit., 10, 1916, 268.) Listed as synonymous with Actinomyces micro- parva Krainsky, 1914 which may be in- tended for Actinomyces microflavvs Kra- insky, Cent. f. Bakt., II Abt., 41, 1914, 662. Nocardia orangica (Berestneff) Chal- mers and Christopherson. {Streptothrix orangica Berestneff, Inaug. Diss., Mos- cow, 1897, quoted from Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 271; Chalmers and Christopherson, idem.) Nocardia rogersi de Mello. (Nocardia (Cohnistreptothrix) rogersi de Mello, A Med. Contemp., 1919; Discomyces rogersi Neveu-Lemaire, Precis Parasitol. Hum., 5th ed., 1921, 44; Actinomyces rogersii Brumpt, Precis de Parasitol., 4th ed., 1927, 1206.) From sputum. Nocardia rubea Chalmers and Chris- topherson. (Ann. Trop. Med. and Para- sit., 10, 1916, 271.) Nomen Jiudum. According to Dodge (Medical Mycology, St. Louis, 1935, 765), this is a synonym of Oospora rubra Wilbert, Recueil Hyg. M^d. V6t. Militaire, 1908. Nocardia saprophytica Chalmers and Christopherson. (Streptothrix leucea saprophytica Foulerton, 1902, quoted from Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Chalmers and Christopherson, idem.) Nocardia urinaria Pijper. (Pijper, 1918, quoted from Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 1057; Actinomyces urinarius Nannizzi, in Pollacci, Tratt. Micopat. Umana, 4, 1934,50.) Oospora hoffmanni (Gruber) Sauva- geau and Radais. (Mikromyces hof- manni (sic) Gruber, Trans. Int. Congr. Hj-g. Derm., VI, 2, 1891-1892, 65 and Arch. f. Hyg., 16, 1893, 35;Sauvageau and Radais, Ann. Inst. Past., 6, 1892, 251; Actinomyces hoffmanni Gasperini, Cent. f. Bakt., 15, 1894, 684; Streptothrix hofmanni Kruse, in Flugge, Die Mikro- organismen, 3 Aufl., 2, 1896, 62; Clado- thrix hoffmanni Mac6, Traitd Pratique de Bact., 4th ed., 1901, 1081; Nocardia hoffmanni Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 268.) From a sample of vaccine. Oospora spumalis Sartory. (Sartory, in Sartory and Bailly, Mycoses pulmo- naires, 1923, 318; Actinomyces spu?7ialis Dodge, Medical Mycology, St. Louis, 1935, 751.) From human sputum. Streptothrix aaser Johan-Olsen. (In- aug. Diss., Christiania, 1893, 91; quoted from Neukirch, Ueber Actinomyceten, Strassburg, 1902, 69.) Streptothrix alpha Price-Jones. (Price- Jones, 1900; quoted from Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270.) Considered synonymous with Streptothrix alba (Rossi Doria). Streptothrix aquatilis Johan-Olsen. (Inaug. Diss., 1893, 93; quoted from Johan-Olsen, Cent. f. Bakt., II Abt., 3, 1897, 279.) Streptothrix beta Price-Jones. (Price- Jones, 1900; quoted from Chalmers and Christopherson, Ann. Trop. Med. and Parasit., 10, 1916, 270; Nocardia beta Chalmers and Christopherson, idem.) Streptothrix chondri Johan-Olsen. (Inaug. Diss., 1893, 95; quoted from Johan-Olsen, Cent. f. Bakt., II Abt., 5, 1897, 278.) Streptothrix enteritidis Pottien. (Quoted from Sanfelice, Cent. f. Bakt., I Abt., Orig., 36, 1904, 355.) FAMILY STREPTOMYCETACEAE 977 Streptothrix foersteri Gasperini. (Gas- perini, Annales de Micrographie, 2, 1890, 462; not Streptothrix foersteri Cohn, Beitr. z. Biol. d. Pflanzen, 1, Heft 3, 1875, 196; Actinomyces saprophyticus Gasperini, Ann. d. 1st. d'Ig. sper. d. Univ. Roma, 2, 1892, 226; Actinomyces saprophyticus var. cromogemis Gasperini, ibid., 229.) From air. Streptothrix gelatinosus Johan-Olsen. (Cent. f. Bakt., II Abt., 3, 1897, 279.) Streptothrix hiimifica Johan - Olsen. (Cent. f. Bakt., II Abt., 3, 1897, 278.) Streptothrix lemani Johan-Olsen. (Inaug. Diss., 1893, 96; quoted from Johan-Olsen, Cent. f. Bakt., II Abt., 3, 1897, 279.) Streptothrix necrophora Wilhelm. (Monats. f. prakt. Tierheilk., H, 1902, 193.) See page 578. Streptothrix leucea Foulerton. (In All- butt and Rolleston, Syst. of Med., 2, 1912, 310.) Streptothrix melanotica Price-Jones- (On the General Characteristics and Pathogenic Action of the Genus Strepto- thrix, 1901 ; also see Foulerton, in Allbutt and Rolleston, Syst. of Med., 2, 1912, 304.) Streptothrix oidicformis Johan-Olsen. (Inaug. Diss., 1893, 96; quoted from Xeukirch, Ueber Actinomyceten, Strass- burg, 1902, 69.) Streptothrix spirilloides Johan-Olsen. (Inaug. Diss., 1893, 96; quoted from Neukirch, Ueber Actinomyceten, Strass- burg, 1902, 69.) Streptothrix tartari Sanfelice. (Cent, f. Bakt., I Abt., Orig., 36, 1904, 355.) Streptothrix xoallemia Johan-Olsen. (Inaug. Diss., 1893, 96; quoted from Neukirch, Ueber Actinomyceten, Strass- burg, 1902, 69.) Streptothrix zopji Casagrandi. (Quoted from Caminiti, Cent. f. Bakt., I Abt., Orig., U, 1907, 198.) Drechsler (Botan. Gazette, 67, 1919, 65 and 147) described eighteen mor- phological types of Actinomyces (Strepto- myces). The relationships of these types to species previously described in the literature are not explained except in four instances. Actinomyces III is re- garded as Actinomyces lavendulae Waks- man and Curtis ; Actinomyces X is re- garded as Streptothrix alba Rossi Doria (possibly Actinomi/ces griseus Krainsky ) ; Actinomyces XII is regarded as Actino- myces aureus Waksman and Curtis; and Actinomyces XVII is Actinomyces scabies Giissow. 978 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus II. Micromonospora ^rskov. (0rskov, Investigations into the morphology of the ray fungi. Copenhagen, 1923, 147; inchides Thermoactinomyces Tsilinsky, Ann. Inst. Past., 13, 1899, 501; ibid., 17, 1903, 206.) Well developed, fine, non-septate mycelium, 0.3 to 0.6 micron in diameter. Grow well into the substrate. Not forming at any time a true aerial mycelium. Multiply by means of conidia, produced singly at end of special conidiophores, on surface of substrate mycelium. Conidiophores short and either simple, branched or produced in clusters. Strongly proteolytic and diastatic. Many are thermophilic and can grow at 65°C. Usually saprophytes. These organisms occur mostly in hot composted manure, dust, soil and in lake bottoms. The type species is Micromonospora chalcca (Foulerton) 0rskov. Key to the species of genus Micromonospora {0rskov Group III). I. Vigorously growing organisms, typically with copious spore formation on glucose- asparagine-agar. A. Vegetative mycelium pale pink to deep orange, no typical soluble pigment. 1. Micromonospora chalcea. B. Vegetative mycelium orange changing to brownish-black, brown soluble pigment. 2. Micromonospora fusca. II. Slowly and feebly growing organisms, with scant spore formation on glucose- asparagine-agar, no soluble pigment. A. Vegetative mycelium pale pink to pale orange. 3. Micromonospora parva. B. Vegetative mj^celium yellow to orange-red. 4. Micromonospora globosa. C. Vegetative mycelium blue. 5. Micromonospora vulgaris. Note : This genus could be subdivided on the basis of the relations of the organisms to temperature, since it includes a number of thermophilic forms which grow readily at 55° to 65°C, mesophilic forms having their optimum temperature at 30°C, and organisms growing at low temperatures in lakes. Each of these can be divided into 3 groups, based on the structure of the spore-bearing hyphae. Among the thermo- philic forms, only representatives of the first group have so far been isolated in pure culture although the existence of the other two groups has definitely been demon- strated in microscopic preparations. These are: Group 1. Simple spore-bearing h^yphae. Group 2. Branching spore-bearing hyphae. Group 3. Spore-bearing hyphae in clusters. 1. Micromonospora chalcea (Fouler- Bact., 1927, 221.) From Greek chalceus, ton) 0rskov. (Streptothrix chalcea Foul- bronze. erton, Lancet, 1, 1905, 1200; Nocardia Description from Jensen, Proc. Linn. chalcea Chalmers and Christopherson, Soc. New So. Wales, 57, 1932, 173. Ann. Trop. Med. and Parasit., 10, 1916, Formation of a unicellular mycelium 268; 0rskov, Thesis, Copenhagen, 1923, which forms distally placed, singly situ- 156; Actiiioinj/ccs chalcca Ford, Tcxtb. of ated spores. No aerial hyphae. No sur- FAMILY STREPTOMYCETACEAE 979 face growth in liquid medium. The organism resists desiccation for at least 8 months. Comparison between the power of resistance of the mycelium and the spores, respectively, will no doubt present great difficulty, because it is al- most impossible to ensure that the two constituents are actually detached. Otherwise, the mj'celium is but slightly capable of germinating, which maj^ be ascertained by inoculating a water agar plate liberall}' with a mixture of mycelial threads and spores. While practically all the spores germinate, the mycelial threads were never found to form new colonies . Vegetative mj^celium on glucose-as- paragine-agar : Heavy, compact, raised, pale pink to deep orange, not spreading much into the medium. Spore-layer well developed, moist and glistening, brown- ish-black to greenish-black, this color sometimes spreading through the whole mass of growth. Gelatin is liquefied. Grows in liquid media as small firm orange granules or flakes. ^lilk is digested with a faintly acid reaction, mostly after a previous coagula- tion. Many strains invert sucros^e. Some strains produce nitrites from nitrates. Starch is hydrolj'zed. Most strains decompose cellulose. Proteolytic actionseems stronger in this than in the other species of this genus. Optimum temperature for growth 30° to 35°C. Thermal death point of my- celium, 70°C in 2 to 5 minutes. Spores resist 80°C for 1 to 5 minutes. Habitat : Soil, lake mud and other sub- strates. In addition to the above ref- erences, see Erikson (Jour. Bact., 4.I, 1941, 299) and Umbreit and :\IcCoy (X Symposium on Hydrobiology, Univ. of Wisconsin Press, 1941, 106-114"). 2. Micromonospora fusca Jensen. (Proc. Linn. Soc. Xew So. Wales, 57, 1932, 178.) From Latin fuscus, dark. Vegetative m3'celium on glucose-as- paragine-agar heavy, compact, orange, rapidly changing to deep brown and nearly black; spore-layer moist, glisten- ing, grayish- to brownish-black. Deep brown soluble pigment. Gelatin is liquefied. Grows in liquid media as small brown granules and flakes. Milk is slowly digested ; no coagulation. Sucrose is inverted. Reduction of nitrates, positive or nega- tive . Cellulose is attacked to a slight extent. Starch is hydrolyzed. Habitat: Soil. 3. Micromonospora parva Jensen. (Proc. Linn. Soc. Xew So. Wales, 57, 1932, 177.) From Latin parvus, small. Scant growth on glucose-asparagine- agar; vegetative mycelium thin, spread- ing widel}^ into the agar, almost colorless to pale pink or orange. Sporulation scant, giving rise to thin grayish, moist crusts on the surface. Gelatin is liquefied. Milk is left unchanged; or coagulated, slowly redissolved with faintly acid reac- tion. Sucrose not inverted. Nitrates not reduced. Cellulose not decomposed. Starch is hydrolyzed. Habitat: Soil. 4. Micromonospora globosa Krassilni- kov. (Ray Fungi and Related Or- ganisms. Izd. Acad. Xauk, Moskow, 1938, 1.34; Microbiology, U. S. S. R., 8, 1939, 179.) From Latin globosus, spheri- cal. A fine (0.5 to 0.8 micron in diameter) monopodially branching mycelium. This mycelium breaks soon into separate pieces of varying length and irregular outline. Conidia are formed at the ends of short branches, one on each. Indi- vidual branches with conidia resemble grape vines. The conidia are spherical 1.0 to 1.3 microns; thej- arise by the 980 MANUAL OF DETERMINATIVE BACTERIOLOGY swelling of the branch tips. The swell- ings become round, acquire the shape of spheres, which, as the formation of the conidia proceeds, are divided from the branch by a transverse septum. Gelatin is liquefied. Colonies : Rugose, at first very com- pact, later acquire a pasty consistency, and their bond with the medium becomes not so fast. The color of the cultures varies from light yellow to orange -red. During fruit-bearing the colonies are covered with a brownish-black tarnish of conidia. In meat-peptone broth, ammonia is produced. Milk: Coagulation; peptonization. Nitrites are produced from nitrates. Sucrose is inverted. Cellulose not decomposed. Starch is hydrolyzed. Habitat: Soil. 5. Micromonospora vulgaris (Tsilin- sky) Waksman, Umbreit and Gordon. (Thermophile Cladothrix, Kedzior, Arch. Hyg., 27, 1896, 328; Thermoaclinonujces vulgaris Tsilinsky, Ann. Inst. Past., 13, 1899, 501 ; Actinomyces rnonosporus Schlitze, Arch. f. Hyg., 67, 1908, 50 (No- cardia monospora Chalmers and Chris- topherson, Ann. Trop. Med. and Para- sit., 10, 1916, 271); Actinomyces glaucus Lehmann and Schiitze, in Lehmann and Neumann, Bakt. Diag., 5 Aufl., 2, 1912, 641 {Nocardia glauca Chalmers and Christopherson, loc. cit.); Micromonos- pora coerulea Jensen, Proc. Linn. Soc. New So. Wales, 57, 1932, 177; Waksman, Umbreit and Gordon, Soil Sci., 47, 1939, 51.) From Latin vulgaris, common. Morphologically the development of this organism is entirely comparable to that of the mesophilic form described by Jensen. The young mycelium shows slightly more branching than that pro- duced by species of Streptomyces. Spores are borne at the end of short branches from which they are easily broken. The aerial mycelium, though present, is usually rudimentarj^, rarely exhibiting the tangled network of strands typical of species of Streptomyces. Ther- mophilic strains of Micromonospora vulgaris differ thus from the mesophilic forms, which show no trace of aerial my- celium. Fragmentation has not been seen in slide cultures of the organism thus far isolated, but it was found to oc- cur in smear preparation. According to Jensen, the mesophilic strains grow slowly on glucose-aspara- gine-agar; vegetative mycelium dense, dark greenish-blue, with a hard and glossy surface. Sporulation very scant. The surface sometimes shows a thin white veil resembling aerial mycelium, but without aerial spores. Gelatin: Liquefaction. Good growth on beef-peptone agar, potato, milk, beef-peptone broth, etc. Grows in liquid media as fairly large, firm, round, white to pink granules (Jensen). Usually a white, powdery, thin aerial mycelium is produced which is hardly raised above the surface. No soluble pigment is formed. Czapek's agar : Growth white, powdery, slightly raised. Broth : A tough white pellicle and in many instances a considerable number of ball-like colonies at the bottom of the tube. No turbidity. Milk: Coagulated and digested. Nitrites not produced from nitrates. Sucrose not inverted. Cellulose not decomposed. Starch is hydrolyzed. Optimum temperature of thermophilic forms 57 °C. Growth range 48° to 68°C. Habitat: Straw, soil, high tempera- ture composts. Appendix: The following anaerobic species has been described : Micromonospora propionici Hungate. (Abst. in Jour. Baot., 48, 1944, 380 and 499; Jour. Bact., 51, 1946, 51.) From the alimentary tract of the wood-eating termite {Amilennes minimus). Fer- ments glucose or cellulose to form acetic and propionic acids and COo. Obligate anaerobe. ORDEE CHLAMYDOBACTERIALES 981 ORDER III. CHLAMYDOBACTERIALES BUCHANAN. (Jour. Bact., 2, 1917, 162.) Filamentous, colorless, alga-like bacteria. May or may not be ensheathed. They may be unbranched or may show false branching. False branching arises from a lateral displacement of the cells of the filament within the sheath which gives rise to a new filament, so that the sheath is branched while the filaments are separate. The sheath may be composed entirely of iron hydroxide, or of an organic matrix impregnated ^^'\\\\ iron, or may be entirely organic. The filaments themselves may show motility by a gliding movement like that found in the blue-green algae (OsciUa- loriaceae). Conidia and motile flagellate swarm cells may be developed, but never endospores. Fresh water and marine forms. Key to the families of order Chlamydobacteriales.* I. Alga-like filaments which do not contain sulfur globules. False branching may occur. A. Usually free floating filaments. ^Motile swarm cells may be formed. Family I. Chlamydohaeteriaceae, p. 981. B. Attached filaments which show a differentiation of base and tip. Xon-molile conidia formed in the swollen tips of the filaments. Family II. Crenothrichaceae , p. 987. II. Alga-like, unbranching filaments which maj- contain sulfur globules when growing in the presence of sulfides. Filaments may be motile by a creeping or sliding movement along a solid substrate. Family III. Bcggiatoaceae, p. 988. FAMILY I. CHLAMYDOBACTERIACEAE MIGULA.** (Arb. Bakt. Inst. Hochschule, Karlsruhe, 1, 1894, 237.) Filamentous bacteria which frequently show false branching. Sheaths may or may not be impregnated with ferric h3'droxide. Cells divide only transversely. Swarm cells, it developed, are usualh' motile by means of flagella. Usually found in fresh water. Key to the genera of family Chlamydobacteriaceae. I. Showing typical false branching. A. Sheaths entirely organic, not impregnated with ferric hydroxide. Genus I. Sphaerotilus, p. 982. B. Sheaths impregnated with ferric hydroxide. Genus II. Clonothrix, p. 983. II. Unbranched or rarely showing false branching. A. Sheaths or holdfasts impregnated with ferric hydro.xide. Genus III. Leptothrix, p. 983. * In Appendix I, p. 996, will be found a group of non-filamentous, non-sheath- forming, colorless sulfur bacteria, as the family Achromatiaceae . Their true rela- tionships are as yet obscure, and they have been attached as an Appendix to the Chlamydobacteriales largeh' on account of the similarity of their metabolism to that of the Bcggiatoaceae. ** Completely revised by Prof. A. T. Henrici, University of Minnesota, Minneap- olis, Minnesota, December, 1938; further revision by Prof. Robert S. Breed, Xew York State Experiment Station, Geneva, New York, July, 1946. 982 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus I. Sphaerotilus KiUzimj. (Kiitzing, Linnaea, 8, 1833, 385; Cladothrix Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 3, 1875, 185.) From Greek sphaera, sphere. Attached, colorless threads, showing false branching, though this may be rare in some species. Filaments consist of rod-shaped or ellipsoidal cells, surrounded by a firm sheath. Multiplication occurs both by non-motile conidia and by motile swarm cells, the latter with lophotrichous flagella. The type species is Sphaerotilus natans Kiitzing. 1. Sphaerotilus natans Kiitzing. (Kiitzing, Linnaea, t, 1926, 34, PI. IV, Figs. 42-43, also var. velatum Plate IV, Figs. 44-46. 9 . Polyangium aureum Krzemieniewski . (Acta Soc. Bot. Pol., 7, 1930, 255.) Etjmiology: Latin, aureus, golden. Separated from Polyangium morula on basis of pigmentation. Fruiting body: Cysts reddish-brown, variable in number, embedded in yellow slime to form a sorus with a common slime envelope. Cysts nearly spherical or slightly elongate, averaging 32 by 37 microns. Cyst wall orange-yellow, about 3.0 microns thick. Older cysts contain shortened rods, a granular mass, and a colorless or yellowish oleaginous liquid. Spores : Rod-shaped. Vegetative cells: Straight rods, of uniform diameter, with rounded ends, 0.7 to 0.9 by 2.8 to 5.3 microns. Habitat: Soil. Illustrations : Krzemieniewski {loc. cit.) Plate XVII, Figs. 14-17. 10. Polyangium stellatum Kofler. (Sitzber. d. kais. Akad. Wiss. Wien, Math. -Nat. Klasse, 122 Abt., 1913, 19.) Etymology: Latin stellatus, stellate. Swarm stage (pseudo Plasmodium) :Not described. Fruiting bodies : Cysts elongate, 80 to 120 microns broad, 160 to 200 microns long, flesh-colored when young, brownish- red when old, star-shaped with 2 to 9 rays fixed by the narrowed base upon a kind of hypothallus. Source and habitat : Found by Kofler (1913, loc. cit.) on hare dung at Vienna. Illustrations: Kofler (1913, loc. cit.) PI.?, Fig. 6. 11. Polyangium ferrugineum Krzemie- niewski. (Acta Soc. Bot. Poloniae, 5, 1927,97.) Etymology : Latin ferrugineus, dark red, like iron rust. Swarm stage (pseudo Plasmodium) :Not described. Fruiting bodies : Irregular, branched and occasionally constricted coils. Branches of same diameter as the main tube. Cyst wall is brown-red. In the interior no differentiation is visible. Rods in cysts are relatively short and thick, 0.8 to 1.1 microns by 2 to 2.5 mi- crons, not definitely arranged. Close to Archangium gephyra, but with cyst walls. Source and habitat : Krzemieniewski {loc. cit.) from soil in Poland and on rabbit dung. Illustrations : Krzemieniewski {loc. cit.) PI. V, Fig. 21. 12. Polyangium indivisum Krzemien- iewski. (Acta Soc. Bot. Poloniae, 5, 1927, 97.) Etymology : Latin indivisus, undivided. Swarm stage (pseudo Plasmodium) : Not described. Fruiting bodies : Similar to Poly- angium Jerrugineum, but much smaller and bright orange-yellow. Enclosed in a similarly colored slime membrane. Interior of coils undifferentiated. Cyst rods 0.8 to 1.0 by 3 to 6 microns, straight, and rounded on ends. Arranged per- pendicularly to the wall, giving a netted appearance resembling Melittangium. Source and habitat : From soils in Poland, Krzemieniewski (1927, loc. cit.). 13. Polyangium spumosum Krzemien- iewski. (Acta Soc. Bot. Pol., 7, 1930, 254.) Etymology : Latin spumosus, foaming, full of foam. 1082 MAXUAL OF DETERMIXATIVE BACTERIOLOGY Fruiting body : Colorless sori embedded in hyaline slime forming a common en- velope around the cysts. Surface white, foamy in appearance; cysts in irregularly rounded accumulations, 100 to ISOmicrons in diameter. Cysts usually spherical, sometimes elongate; 18 to 38 by 20 to 50 microns; average 28 by 34 microns. Cyst membrane colorless. Cysts con- tain bundles of shortened cells, a granu- lar colorless mass, and a clear oleaginous fluid. Spores: Shortened rods. Vegetative cells: Straight rods, uni- formly thick, with rounded ends; 0.6 to 0.8 by 3.9 to 6.8 microns. Habitat: Soil. Illustrations: Krzemieniewski {loc. cit.) Plates XVI-XVII, Figs. 10-13. 14. Polyangium fumosum Krzemien- iewski. (Acta Soc. Bol. Pol., 7, 1930, 253.) Etymology : Latin fumosus, smoky. Fruiting body: A flat, crust-like layer of 2 to 20 (or more) cysts arranged to form a sorus. Sori rounded, up to 90 microns in diameter, or irregularly shaped; often elongate up to 400 microns long. Smoky- gray color due to surrounding slime walls. Outer profile of sheath (or cortex) ir- regular. Cyst wall 2.4 to 3.5 microns thick; cysts often nearly spherical, 13 to 48 microns in diameter, though fre- quently elongate. Average 36 by 44 microns. Colorless, single, inclosed in a transparent membrane. Spores : No data. Vegetative cells: Long, straight, cy- lindrical with rounded ends; 0.7 to 0.9 by 2.7 to 5.7 microns. Encysted cells similar. Habitat: Soil. Illustrations : Krzemieniewski (loc. cit.) Plate XVI, Figs. 6-9. 15. Polyangium parasiticum Geitler. (Arch. f. Protistenkunde, 50, 1924, 67.) Etymology : Latin parasiticus, para- sitic. Swarm stage (pseudo Plasmodium) : In water, on surface of the alga Cladophora. Pseudoplasmodia small. Rods long, cy- lindric, rounded at end and 0.7 by 4 to 7 microns. At first saprophytic, later entering and destroying the Cladophora cell. Fruiting bodies : Sometimes single, usually 2 to 8 microscopically small, united in irregular masses, spherical ori somewhat elongated. From 15 to 50 microns, usually 25 to 40 microns, with hyaline slime. When mature, red-brown in color, with firm wall. Source and habitat : Found on Clado- phora (fracta?) in pool at Vienna (Geitler, 1924). Illustrations: Geitler (1924, loc. cit.) Figs. 1-10. Genus II. Synangium Jahn. (Jahn, Beitrage zur botan. Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 79; Apelmocoena Enderlein, Bakterien-Cyclogenie, Berlin, 1924, 243.) Etymology: Greek syn, together and angion, vessel, referring to the clustering of the cysts. Diagnosis: Cysts provided with an apical point, united more or less completely to rosette-shaped, hemispherical or spherical fruiting bodies. The type species is Synangium sessile (Thaxter) Jahn. Key to the species of genus Synangium. I. Cysts irregular, pointed, united as a rosette on a slimy base, without a stalk. 1. Synangium sessile. II. The fused cysts on a simple or branched stalk. A. Cyst group spherical, with the points of the cysts covered as with hair, reddish. 2. Synangium lanuginosum. FAMILY POLYANGIACEAE 1033 B. Cyst group an oblate spheroid, yellow. Points of cysts less numerous. 3. Synangium thaxteri. 1. Synangium sessile (Thaxter) Jahn. {Chondromyces sessilis Thaxter, Bot. Gaz., 37, 1904, 411; Jahn, Beitnige zur botanischen Protistologie. I. Die Poly- angiden, Geb. Borntraeger, Leipzig, 1924, 79.) Etymology: Latin sessilis, sessile, not stalked. Swarm stage (pseudoplasmodium) :Not described. Fruiting body : Cysts form on the base a clump or rosette without trace of stalk. Diameter of rosettes 100 to 250 micron.s. Individually the cysts are quite variable in form, irregularly spindle-shaped, usu- ally short-pointed, wrinkled surface toward the tip. At the base they fuse or unite to irregular masses. Cysts 18 to 55 by 25 to 75 microns, average 40 by 50 microns. Source and habitat : Thaxter (loc. cil.) found this on decaj'ing wood in Florida. Illustration: Thaxter {loc. cit.) PL 27, Figs. 14-15. 2. Synangium lanuginosiim (Kofier) Jahn. {Chondromyces lanuginosus Kofier, Sitzber. d. Kais. Akad. Wiss. Wien. Math. -Nat. Klasse, 122 Abt., 1913, 861; Jahn, Beitrage zur botanischen Protis- tologie. I. Polyangiden, Geb. Born- traeger, Leipzig, 1924, 79.) Etymology : Latin lanuginosus, woolly. Swarm stage (pseudoplasmodium) :Not described. Fruiting body : Cyst cluster, consisting of united cysts, spherical or oval, 80 to 200 microns in diameter, when dry, dark flesh-colored, covered with hairs 15 to 50 microns long, originating from the indi- vidual cysts and giving the cyst cluster the appearance of a hairy ball. Skin of the cysts not definite. Rods within the cysts 3 to 6 microns. The cyst clusters are terminal on more or less forked stalks, about 1 mm high. Source and habitat: Kofler {loc. cit.) found this on rabbit dung at Vienna. Illustrations: Kofier {loc. cit.) PI. 1, Figs. 1-3. 3. Synangium thaxteri (FauU) Jahn. {Chondromyces thaxteri Faull, Bot. Gaz., 62, 1916, 226; Jahn, Beitrage zur botan- ischen Protistologie, I. Die Polyangiden. Geb. Borntraeger, Leipzig, 1924, 79.) Regarded as a synonj^'m of Synangium lanuginosum bj- Krzemieniewski, Acta Soc. Bot. Poloniae, 4, 1926, 39. Etymology : Named for Dr. Roland Thaxter, American botanist. Swarm stage (pseudoplasmodium): Cultured for 2 years on dung, best in mixed cultures. Rods 0.5 by 3 to 6 microns. Fruiting body : Fruit cluster flattened, spherical, yellow to flesh color or red- dish-orange, with a stalk which varys in length, about 140 microns in diameter. The bristles corresponding to the single cysts are 15 to 30 microns long, at the base 10 to 12 microns wide. Sometimes cyst single, usually 3 to 4, occasionally 20 to 30. Rods 0.5 by 3 to 6 microns. Stalk maximum length 0.75 mm, usually 350 microns, single or branched. Broad based, narrowing to apex and yellow in color. In germination rods move from basal scar of membrane, leaving the empty sack behind. Source and habitat : On deer dung in Ontario, Canada (Faull). Illustrations: Faull {loc. cit.) PI. 5 and 6. Jahn {loc. cit.) Fig. X, p. 80. Genus III. Melittangium Jahn. (Jahn, Beitrage zur botan. Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 78.) Etymology: Greek melitta, bee and angion, vessel, because of the honey -comb pattern of the membrane. 1034 MANUAL OF DETERMINATIVE BACTERIOLOGY Diagnosis : Cysts brownish orange-red, on short white stalk, like a mushroom. Has appearance of a white-stalked Boletus. The rods inside stand at right angles to the membrane. Upon germination the covering membrane is left colorless and with an appearance of honey -comb. The type species is Melittangium boletus Jahn. 1. Melittangixun boletus Jahn. (Bei- trage zur botanischen Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 78.) Etymology: Latin boletus, a kind of mushroom . Swarm stage (pseudoplasmodium) : Xo description. Fruiting bodies : Cyst stalked, mush- room-like, white when immature, then yellowish-flesh colored, finally yellowish- brown to nut brown, when dried more reddish-brown. Larger diameter of cyst about 100 microns, height 40 to 50 microns, length of white stalk about 40 microns, length of rods in the cyst 3 to 4 microns by 0.5 microns. Sometimes the cyst is smaller and spherical (50 to 60 microns diameter), sometimes there is fusion of neighboring cysts, occasionally the stalk is abortive. Source and habitat: Jahn {loc. cit.) found this not uncommon on rabbit and deer dung in the vicinity of Berlin, also on deer dung from Denmark. Krzemien- iewski (1927, loc. cit.) reported it as common in Polish soils. Illustrations: Jahn {loc. cit.), PI. 2, Fig. 17 and 18. Also Fig B, p. 11, C-F, p. 23, 0-Q, p. 43, T-U,p. 55. Krzemien- iewski, Acta Soc. Bot. Poloniae, 4, 1926, 1, PI. V, Fig. 55-56. Genus IV. Podangium Jahn. (Cystobacter Schroeter, in Cohn, Kryptogamenflora v. Schlesien, 3, 1, 1886, 170; Jahn, Beitriige zur botan. Protistologie. I. Die Polyangiden. Geb. Borntraeger, Leipzig, 1924, 80; Monocystia Enderlein, Bakterien-Cyclogenie, Berlin, 1924, 243.) Etymology: Greek pus, podis, foot and angion, vessel. Diagnosis: Cysts chestnut -brown or. red-brown, single on a more or less definite white stalk. The type species is Podangium erectum (Schroeter) Jahn. Key to the species of genus Podangium. I. Stalk scarcely definite, cysts short, appressed, if elongate then passing over from the white stem into the club-shaped cyst. Ripe cysts chestnut-brown. 1. Podangium erectum. II. Stalk well differentiated. A. Cysts spherical, often irregular, confluent, the white stalk short. 2. Podangium lichenicolum . B. Cysts lengthened ellipsoidal, red-brov\-n, definitely differentiated from the white, slender stalks. 3. Podangium gracilipes. 1. Podangium erectum (Schroeter) Jahn. {Cystobacter erectus Schroeter, in Cohn, Krytogamenflora v. Schlesien, 3, 1, 1886, 170; Chondromyces erectxis Thax- zur botanischen Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 80.) Etymology: Latin erectus, erect, up- ter, Bot . Gaz .,23, 1897 , 407 ; Jahn , Beitriige right . FAMILY POLYANGIACEAE 1035 Swami stage (pseudo Plasmodium) : Kofler states rods are 2 to 5 microns in length. Fruiting bodies : Cysts usually short, almost spherical, compact, rounded above, orange-red changing to chestnut-brown, single on a white to yellow hypothallus constituted from the slime remaining behind. A definite "foot" of whitish slime is seldom observed. Fifty to hundreds together. Usually about 80 microns high and 40 to 50 microns broad above, smaller below, often spherical cysts 60 microns in diameter. Rods in cysts 0.6 by 4 microns. Jahn believes the European form to be distinct from that described by Thaxter. Thaxter's form produces cystophores 60 to 300 microns long which wither at maturity so that cysts appear sessile. Source and habitat : Thaxter {loc. cit.), horse dung in laboratory cultures, Massa- chusetts. Kofler (Sitzber. d. Kais. Akad. Wiss. Wien. Math. -Nat. Klasse, 122 Abt., 1913), mouse dung. Jahn (1924) common on manure of different kinds, also on bark covered with lichens. Krzemieniewski (Acta Soc. Bot. Pol., 5, 1927, 102) re- ported this species from Polish soil, but rare. Illustrations: Thaxter (loc. cit.) PI. 31, Figs. 16-19. Quehl, Cent. f. Bakt., II Abt., 16, 1906, PI. 1, Figs. 4. Jahn {loc. cit.) PI. I, Figs. 7, 8, and 9. Krze- mieniewski, Acta Soc. Bot. Poloniae, 4, 1926, 1, PI. V, Figs. 52-53. 2. Podangium lichenicolum (Thaxter) Jahn. {Chondromyces lichenicolus Thax- ter, Bot. Gaz., 17, 1892, 402; Jahn, Beitrage zur botanischen Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 81.) Etymology: Greek lichen, tree moss, lichen. Latin -colus, dwelling. Swarm stage : Reddish, rods cylin- drical, tapering slightly, 0.6 by 5 to 7 microns. Germinate readily after dry- ing for 18 months when sown on moist lichens. Fruiting bodies : Cysts single, rounded or irregularlj' lobed, often confluent. Cj'stophore short, squarish, often lacking or misshapen. Cysts 28 to 35 microns, stem 7 to 8 by 10 microns. Source and habitat: Thaxter (1892), parasitic upon living lichens, which it destroys, New Haven, Conn. Thax- ter (1904, loc. cit.), lichens, Indiana, on algae, seen on wet boards, in mill race, Massachusetts. Illustrations: Thaxter (1892, loc. cit.) PI. 23, Figs. 20 to 23. Quehl, Cent. f. Bakt., II Abt., 16, 1906, 9, PI. 1, Fig. 6. 3. Podangium gracilipes (Thaxter) Jahn. {Chondromyces gracilipes Thax- ter, Bot. Gaz., 23, 1897, 406; Jahn, Bei- trage zur botanischen Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leip- zig, 1924,82.) Etymology: Latin gracilipes, slender footed. Swarm stage: Rods 5 to 7 microns. Fruiting bodies : Cysts bright orange- red, or red, 25 by 35 microns, elongate, rounded, on a white pointed stalk, rigid and persistent on substratum, rods also in stalk. Shortened rods in cyst 3 to 5 microns. Cysts sometimes pear-shaped, caducous. Source and habitat : Thaxter (loc. cit.), from rabbit dung, Massachusets. Kofler (1913, loc. cit.), dung, Vienna. Jahn (loc. cit.) relatively common. Twice on rabbit dung near Berlin, once on goat dung in Norwa3^ Krzemieniewski (1927 loc. cit.) reported this species from Polish soil, but rare. Illustrations : Thaxter {loc. cit.) PI. 31, Figs. 20-24. Quehl, Cent. f. Bakt., II Abt., 16, 1906, PI. 1, Fig. 12. Jahn {loc. cit.) PI. II, Figs. 19, 20. Krzemien- iewski (1926, loc. cit.), Pi. V, Fig. 54. 1036 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus V. Chondromyces Berkeley and Curtis. (See Berkeley, Introduction to Cryptogamic Botany, London, 1857, 313; Sligmalella Berkeley and Curtis, ibid. 1857, 313 (figure but no description) ; Berkeley (descrip- tion). Notes on North American Fungi, Grevillea, 3, 1874, 97; see Berkeley and Curtis, in Saccardo, Sylloge Fungorum, 4, 1886, 679, Polycephalum? Kalchbrenner and Cooke, Grevillea, 9, 1880, 22; Myxobolyrs Zukal, Ber. d. deutsch. Bot. Gesellsch., U, 1896, 346; Cysfodesmia P^nderlein, Bakterien-Cyclogenie, Berlin, 1924, 243.) Synonymy : A species was figured and named in 1857 by Berkeley as Chondromyces crocatus Berkeley and Curtis, but not described. The generic name was finally described in 1874. Probably the date of the name should be the date of its description, although it is possible that an adequate labeled illustration should be interpreted as valid publication. Etymology : Greek chondros, grain and myces, (fungus). Diagnosis: Cysts compactly grouped at the end of a colored stalk (cystophore). Cystophore simple or branched. The type species is Chondromyces crocatus Berkeley and Curtis. Key to the species of genus Chondromyces. I. Cysts not in chains. A. Cysts sessile when ripe. 1. Cysts not pointed. a. Cysts rounded, b. Yellow. 1. Chondromyces crocatus. bb. Bright orange -red. 2. Chondromyces aurantiacus. aa. Cysts cylindrical. 3. Chondromyces cylindriciis. 2. Cysts pointed. 4. Chondromyces apiculatus. B. Cysts borne on stalk or stipe when ripe. 1. Cysts orange-colored and truncate or rounded at distal end. a. Cystophore usually simple. 5. Chondromyces pediculatus. aa. Cystophore usually branched. 6. Chondromyces niedius. 2. Cysts copper-red when ripe; pear-shaped. 7. Chondromyces minor. II. Cysts in chains at end of a compact stalk. 8. Chondromyces catenulatus. 1. Chondromyces crocatus Berkeley Etymology: Latin crocatus, saffron and Curtis. (Berkeley, Introduction yellow. to Cryptogamic Botany, London, 1857, Swarm stage (pseudoplasmodium) : 313; Berkeley, Notes on North American Pale orange-red. Rods cylindrical or Fungi, Grevillea, 3, 1874, 64; Mi/xo6o;UAL OF DETERMIXATIVE BACTERIOLOGY I. Die Polyangideu, Geb. Bonitraeger, Leipzig, 1924, 86.) Etymology: Greek megalos, large; sporos, seed, spore; large spored. Swarm stage (pseudoplasmodium) : Not described. Fruiting bodies : About 80 to 160 mi- crons wide, rounded, cushion-shaped, dark flesh color. Spores 2 microns. Source and habitat : Jahn {loc. cit.) , on stag dung near Berlin. Illustrations: Jahn {loc. cit.) Fig. Y, i to k, p. 87. 4. Chondrococcus macrosporus Krze- mieniewski. (Acta Soc. Bot. Poloniae, 4, 1926). According to Krzemienievvski, not to be confused with Zukal's species, Myxococcus macrosporus (Ber. d. deutsch Bot. Gesellsch., 15, 1897, 542.) Etymology: Greek makros, long, large; sporos, seed, spore; large -spored. Swarm stage (pseudoplasmodium) :Not described. Fruiting bodies: Much like Chondro- coccus coralloides, differing in color and in size of spores. Spores 1.6 to 2.0 microns. Fruiting body yellow or light brown color, with long branches. Source and habitat : Krzemieniewski {loc. cit.), found it first on leaves, later isolated from soil on rabbit dung. Illustrations: Krzemieniewski {loc. cit.) PI. II, Fig. 19. 5. Chondrococcus blasticus Beebe. (Iowa State Col. Jour. Sci., 15, 1941, 310.) Etymology : Greek blastikos, budding. Fruiting body : Primary : Spherical to subspherical, usually sessile but occa- sionally with a short stalk or foot; pale pink to bright salmon pink; 300 to 600 microns in diameter. No outer wall or limiting membrane evident. Develops on sterilized rabbit dung in from 3 to 6 days at room temperature. Secondary: Arising as bud-like growth from the primary fruiting body. Develops into irregularly shaped, finger-, coral- or bud- like protuberance. Seldom branched; occasionally stalked but usually sessile on primary fruiting body until latter is utilized in formation of several secondary fruiting bodies. Deep pink to salmon pink in color. Variable in size and shape ; 50 to 150 by 75 to 225 microns. No outer wall or limiting membrane evident. Spores : Spherical, thick-walled, highly retractile; 1.2 to 1.4 microns in diameter. Held together in the fruiting body by the mass of slime. Vegetative cells : Long, slender, flexible rods, straight or curved to bent, ends rounded to slightly tapered, Gram- negative. 0.5 to 0.6 b3'3.0 to 5.0 microns. Usually found in groups of 2 to 12 lying parallel on the surface of the slimy colony, the group moving as a unit. Motile by a crawling or creeping motion, no flagella. Vegetative colony: Thin, colorless, transparent at margin ; surface broken by many small ridges or veins. Center smooth, slightly thicker, often showing pale pink color. Fruiting bodies first form at or near center, later distributed irregularly on other parts of colony. Margin composed of active vegetative cells. Physiology : Good growth on mineral salt agar to which has been added such complex carbohydrates as dulcitol, inulin, cellulose, reprecipitated cellulose or starch; starch hydrolyzed, cellulose not destroyed appreciably. Can utilize agar as both C and N sources. Best growth on suspensions of killed bacterial cells in agar. Growth inhibited partially or entirely by arabinose, mannose and maltose. Source: Goat dung and soil, Ames, Iowa. Habitat: Soil. Decomposes organic matter, especially bacterial cells in dung. Illustrations: Beebe {loc. cit.) PI. II, Figs. 5-6, pi. IV, Fig. 18. 6. Chondrococcus cerebriformis (Kofier) Jahn. {Myxococcus cerebri- formis Kofler, Sitzber. d. kais. Akad. Wiss., Wien. Math. -Nat. Klasse, 122 FAMILY MYXOCOCCACEAE 1047 Abt., 1913, 866; Jahn, Beitriige zur botanischen Protistologie. I. Die Polyan- giden, Geb. Borntraeger, Leipzig, 1924, 86.) Etymology: Latin cerehrinn, brain; formis, shape. Swarm stage (pseudoplasmodium) : Rods 4 to 12 microns. Fruiting bodies: About 1 mm long, clumped masses with swollen upper surface, brain-like, violet rose, often lead- gray. Cysts 100 to 170 microns, without slime envelope. Spores 1.1 to 1.6 mi- crons. Jahn (loc. cit.) suggests that this may be Archangium gephyra. Source and habitat : Kofier {loc. cit.). on hare dung in the vicinity of Vienna. Illustrations: Kofier {loc. cit.) PI. 2, Figs. 7 and 8. 7. Chondrococcus columnaris (Davis) Ordaland Rucker. {Bacillus columnaris Davis, Bull. U. S. Bur. Fisheries, 38, 1923, 261; Ordal and Rucker, Proc. Soc. Exper. Biol, and Med., 56, 1944, 18; also see Fish and Rucker, Trans. Amer. Fish. Soc, 73, 1944 in press; Cyfophaga colum- naris Garnjobst, Jour. Bact., 49, 1945, 113.) Etymology: From Latin columnaris, rising in the form of a pillar. Vegetative cells: Flexible, weakly refractive, Gram-negative rods, 0.5 to 0.7 by 4 to 8 microns. Creeping motion observed on solid media, and flexing movements in liquids. Spores (microcj'sts) : 0.7 to 1.2 microns, spherical to ellipsoidal, occurring on both liquid and solid media. Physiology: Growth best on 0.5 to 0.9 per cent agar with 0.25 to 0.50 per cent Bactotryptone at pH 7.3. Colonies on tryptone agar yellow, flat and irregu- lar. Edge uneven with swarming appar- ent. Gelatin liquefied rapidly. No in- dole. No reduction of nitrates. Starch, cellulose and agar not attacked. Sugars not fermented, but glucose oxidized. Fruiting bodies on agar not deliques- cent, and surrounded by a firm mem- brane. A peculiar type of fruiting body formed in liquid media. Where organ- isms are in contact with infected tissues or with scales, produce columnar, some- times branched, fruiting bodies in which typical spores (microcysts) develop in 7 to 10 days. Source and habitat: First described as cause of bacterial disease of warm water fishes (Davis, loc. cit.) and later in fingerlings of the cold water blue black salmon {Oncorrhynchus nerka). Transmissible to salmonid fishes. Genus III. Angiococcus Jahn. (Beitriige zur Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 89.) A segregate from Myxococcus Thaxter. Diagnosis: Fruiting body consisting of numerous round (disk-shaped) cysts, cyst wall thin, spores within. Etymology: Greek angion, vessel and kokkos, coccus (ball). The type species is Angiococcus disciformis (Thaxter) Jahn. Key to the species of genius Angiococcus. A. Cysts yellow to dark orange-yellow; disk-shaped; 35 microns in diameter. 1. Angiococcus disciformis. B. Cysts colorless to yellow; round; up to 15 microns in diameter. 2. Angiococcus cellulosum. 1. Angiococcus disciformis (Thaxter) Jahn. {Myxococcus disciformis Thaxter, Bot., Gaz., 37, 1904, 412; Jahn, Beitrage zur botanischen Protistologie. I. Die Polyangiden, Geb. Borntraeger, Leipzig, 1924, 89.) 1048 MANUAL OF DETERMINATIVE BACTERIOLOGY Etymology: Greek diskos, a quoit, dis- cus; Latin formis, shape. Swarm stage (pseudoplasmodium ) : Rods 0.5 to 0.6 by 2 to 3 microns. Fruiting bodies: Cysts disk-shaped, crowded, sessile, attached by a more or less ragged scar-like insertion, or in masses. Cysts yellowish when young, when old dark orange-yellow, about 35 by 10 microns. Cyst wall distinct, thin, be- coming very slightly wrinkled. Spores irregularly spherical, embedded in vis- cous slime, difficult to see in the ripe cyst. Source and habitat : Thaxter {loc. cit.), dung of muskrat and deer, Massachusetts and New Hampshire. Krzemieniewski (1927, loc. cit.), rare in Polish soils. Illustrations : Thaxter (loc. cit.) PI. 27, Figs. 19-21. Krzemieniewski Acta Soc. Bot. Poloniae, 4, 1926, PI. II, Figs. 21-22. 2. Angiococcus cellulosum Mishustin. (Microbiology, Moscow, 7, 1938, 427.) Etymology: Modern Latin cellulosum, cellulose. Fruiting body : Regularly rounded (less frequently extended or angular), 20 to 150 microns in diameter; yellow or pink in color, to drabbish when old. Encysted cells surrounded by a colorless cyst wall or envelope. Usually 1 to 3 short stalks or cystophores up to 10 microns high. Within outer wall are numerous cysts containing resting cells (spores). Cysts have regularly rounded form; unpig- mented to yellow; 5 to 15 microns in diameter, average 6 microns. Number of cysts ill fruiting body increases with age. Spores : Cocci (term used is shortened rods) combined into globular aggrega- tions easily broken up. Size not given. Vegetative cells: 0.4 to 0.5 by 1.5 to 2.0 microns. Cell contents pigmented gray, and of indefinite outline (?). Vegetative colony : Fairly rapid growth on cellulose with silica gel . Colony has a yellowish cast. Reaches diameter of 1.5 to 2.0 cm after 6 days with center yellowish-pink and margin tinged light pink. Surface moist. Fruiting bodies more numerous at center, but distributed over entire area. Fruiting bodies do not noticeably protrude above the surface of the colony. Physiology : Cellulose attacked but not completely destroyed. Lower fibers re- main intact, but on treatment with hot soda solution they fall spart. Habitat : Soils. Genus IV. Sporocytophaga Stanier. (Jour. Bact., 40, 1940, 629.) Diagnosis : Spherical or ellipsoidal microcysts formed loosely in masses of slime among the vegetative cells. Fruiting bodies absent. Etymology: Greek sporos seed, spore; kytos hollow place, cell and phagein to eat. The type species is Sporocytophaga myxococcoides (Krzemieniewska) Stanier. Key to the species of genus Sporocytophaga. I. Microcysts spherical. A. Does not utilize starch. B. Utilizes starch. II. Microcysts ellipsoidal. 1. Sporocytophaga myxococcoides. 2. Sporocytophaga congregata. 3. Sporocytophaga ellipsospora. FAMILY MYXOCOCCACEAE 1049 1 . Sporocytophaga myxococcoides (Krzemieniewska Stanier. {Spirochaeta cytophaga Hutchinson and Clayton, Jour. Agr. Sci., 9, 1919, 150; Cyto- phaga myxococcoides Krzemieniewska, Arch. Mikrobiol., 4, 1933, 400; Cytophaga globulosa Stapp and Bortels, Cent. f. Bakt., II Abt., 90, 1934, 47; Cytophaga hutchinsonii Imsenecki and Solntzeva, Bull. Acad. Sci. U.S.S.R., Ser. Biol., Xo. 6, 1936, 1129; not Cytophaga hutchin- soni Winogradsky, Ann. Inst. Pasteur, 4S, 1929, 578; Stanier, Jour. Bact., 40, 1940, 630.) Etymology : Modern Latin from generic name Myxococcus, and eidos, like. Vegetative morphology: Flexible, singly occurring rods, 0.3 to 0.4 micron wide at the center, tapering to both ends. Length 3 to 8 microns according to Krzemieniewska {loc. cit.), 2.5 to 5 microns according to Jensen (Proc. Linn. Soc. X.So. Wales, 65, 1940, 547). May be straight, bent, U-shaped or S-shaped. Show creeping motility (Stapp and Bortels, loc. cit.). Stain poorly with ordinarj^ aniline dyes; with Giemsa's stain, the young cells are colored uni- formly except for the tips. As the rods shorten and swell to form microcysts, the chromatin becomes concentrated and moves toward the center of the cell, generally in the form of two parallel bands (Krzemieniewska, Acta Soc. Bot. Pol., 7, 1930, 514). Microcysts: Spherical, 1.3 to 1.6 microns in diameter, covered with a sheath of mucus. According to Krze- mieniewska (1930, loc. cit.), germination is by emergence of the shortened rod from the sheath, followed by elongation; according to Stapp and Bortels {loc. cit.) and Imsenecki and Solntzeva (loc. cit.), by a simple elongation of the entire microcyst. Growth is strictly confined to cellulose. On mineral salts-silica gel plates covered with filter paper, yellow, glistening, slightly mucilaginous patches are pro- duced after a few days. The color gradually assumes a light brownish tinge on aging. The filter paper in these regions is eventually completely dis- solved and the patches become trans- lucent. Ammonia, nitrate, asparagin, aspartic acid and peptone can serve as sources of nitrogen (Jensen, loc. cit.). Strictly aerobic. Optimum temperature 28 to 30 °C. Source : Isolated from soil . Habitat: Soil. Decomposes cellulose. 2. Sporocytophaga congregata Fuller and Xorman. (Jour. Bact., 4-5, 1943, 567.) Etymolog}' : Latin congrego, to as- semble. Vegetative cells are long, fiexuous rods with pointed ends, 0.5 to 0.7 by 5.5 to 8.0 microns. Creeping motility on solid surfaces. Spores (microcysts) : Spherical, 0.7 to 1.1 microns in diameter. Usually occur in localized regions within the colony. Growth on starch agar is smoky, later turning yellow. Colonies are irregularly round, slightly concave. Edge is smooth and entire at first, later becoming irregu- lar. Marginal and internal swarming may be prominent. The vegetative cells gather into groups and in these regions a large number of spherical spores are found. Growth on cellulose dextrin agar is pale; colonies are small and concave. Hollowing of the agar is limited to the area of colony growth. Glucose, galactose, lactose, maltose, sucrose, arabinose, calcium gluconate, starch, cellulose dextrin, pectin, and hemicellulose are utilized. Filter paper is not attacked. Ammonium, nitrate, and peptone are suitable nitrogen sources. Indole not formed. Xitrites not produced from nitrates. 1050 MANUAL OF DETERMINATIVE BACTERIOLOGY Litmus milk : Growth but no digestion or curd formation. Highly aerobic. Optimum temperature 25° to 30°C. Source : Isolated from soil. Habitat: Soil. Decomposes organic matter. 3. Sporocytophaga ellipsospora (Im- senecki and Solntzeva) Stanicr. (Cjjto- phaga ellipsospora Imsenecki and Solnt- zeva, Bull. Acad. Sci. U.S.S.R., Ser. Biol., No. 6, 1936, 1137; Stanier, Bact. Rev., 6, 1942, 153 and 190.) Etymology: Greek ellipsis, an ellipse, and spores, seed. Vegetative morphology : Flexible, singly occurring rods, 0.4 micron wide at the center and tapering to both ends. Length 7.5 microns. May be straight, bent, U-shaped, or S-shaped. Show creeping motility. Microcysts : Oval or somewhat elon- gated, 0.9 to 1.2 by 1.6 to 1.8 microns Almost always situated in closely-packed aggregates, isolated individual micro- cysts rare. Germinate bj^ elongation. Growth is strictly confined to cellulose. On mineral salts-silica gel plates covered with filter paper, orange, glistening, mucilaginous patches are produced. Ul- timately the filter paper is completely dissolved and the patches become trans- lucent. Ammonia, nitrate and peptone can serve as sources of nitrogen. Strictly aerobic. Optimum temperature 28° to 30°C. Source: Isolated from soil. Habitat : Soil. Decomposes cellulose FAMILY SPIROCHAETACEAE 1051 ORDER V. SPIROCHAETALES BUCHANAN.* (Jour. Bact., 3, 1918, 542.) Slender, flexuous cell body in the form of a spiral with at least one complete turn, 6 to 500 microns in length. Some forms maj^ show an axial filament, a lateral crista or ridge, or transverse striations; otherwise no significant protoplasmic pattern. Smaller forms may have a lower refractive index than bacteria, and so living organisms can be seen only with dark field illumination. Some forms take aniline dj^es with difficulty. Giemsa's stain is uniformly successful. ^lultiplication by transverse fission. Xo sexual cycle known. Granules formed by some species in vector hosts. All forms are motile. Xo organs of locomotion**; motility serpentine or by spinning on the long axis without polaritj'. Free-living, saprophytic and parasitic. Key to the families of order Spirochaetales. I. Spirals 30 to 500 microns in length, having definite protoplasmic structures. Family I. Spirochaetaceae, p. 1051. II. Spirals -4 to 16 microns in length, having no obvious protoplasmic structure. Family II. Treponemataceae, p. 1058. FAMILY I. SPIROCHAETACEAE SWELLENGREBEL. (Ann. Inst. Past., 21, 1907, 581.) Coarse spiral organisms, 30 to 500 microns in length, having definite protoplasmic structures. Found in stagnant, fresh or salt water and in the intestinal tract of bivalve molluscs (Lamellibranchiata) . Key to the genera of family Spirochaetaceae. I. Xo obvious periplast membrane and no cross-striations. Genus I. Spirochaeta, p. 1051. II. Periplast membrane present. Cross-striations prominent in stained specimens. A. Free -living in marine ooze. Genus II. Saprospira, p. 1054. B. Parasitic on lamellibranch molluscs. Crista prominent. Genus III. Cristispira, p. 1055. Genus I. Spirochaeta Ehrenberg. (Ehrenberg, Abhandl. Berl. Akad., 1833, 313; Spirochoeta Dujardin, Hist. nat. des Zoophytes, 1841, 209; Spirochaete Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 1, 1872, * Originally prepared by Prof. D. H. Bergey, Philadelphia, Pa., October, 1922. Revised by Prof. E. G. D. Murray, ]^IcGill University, Montreal, P. Q., Canada, December, 1938. Further revision by Dr. G. H. Robinson, Wm. H. Singer Memorial Research Laboratory of the Allegheny General Hospital, Pittsburgh, Pa., September, 1943. Appendices prepared by Mrs. Eleanore Heist Clise, Geneva, X. Y., Septem- ber, 1943. ** Recent photographs taken with the electron microscope indicate the presence of structures resembling flagella (Mudd, Polevitzky and Anderson, Jour. Bact., 46, 1943, 15). Whether these can be considered as organs of locomotion awaits decision. At present it seems best to confine descriptions of structure to features seen bj'^ use of the oil immersion lens. 1052 MANUAL OF DETERMINATIVE BACTERIOLOGY 180; Ehrenhergia Gieszczykiewicz, Bull. Acad. Polonaise d. Sci. et Lettres, CI. Sci. Math, et Nat., S^r. B, 1939, 24.) Non-parasitic, with flexible, undulating body and with or without flagelliform tapering ends. Protoplast wound spirally around a well-defined axial filament. No obvious periplast membrane and no cross-striations. Motility by a creeping motion. Primary spiral permanent. Free-living in fresh or sea water slime, especially in the presence of H^S. Common in sewage and foul waters. The type species is Spirochaela plicatilis Ehrenberg. Key to the species of genus Spirochaeta. I. Large spirals with rounded ends. II. Smaller spirals with pointed ends. 1. Spirochaeta plicatilis. 2. Spirochaeta marina. 3. Spirochaeta curystrepta. 4. Spirochaeta steiiostrepta. 5. Spirochaeta daxensis. 1. Spirochaeta plicatilis Ehrenberg. (Ehrenberg, Die Infusionstierchen, 1838, 83; Spirillum plicatile Dujardin, Infus., 1, 1841, 225; Spirulina plicatilis Cohn, Nova Acta Acad. Caes. Leop. Carol., 2.^, I, 1853, 125; Spirochaete plicatilis Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 2, 1872, 180; Spirillum portae Mantegazza, Giorn. 1st. Lomb., 3, p. 487, according to DeToni and Trevisan, in Saccardo, Sylloge Fungo- rum, 8, 1889, 1006; Spirochaeta plicatilis plicatilis Zuelzer, Arch. f. Protistenk., 24, 1912, 17; Ehreribergia plicatilis Giesz- czykiewicz, Bull. Acad. Polonaise d. Sci. et Lettres, CI. Sci. Math, et Nat., Ser. B, 1939,24.) From Latin, folded. Cylindrical : 0.5 to 0.75 by 100 to 500 microns, with blunt ends. Spiral amplitude : 2.0 microns, regular. Spiral depth: 1.5 microns, regular. Waves, several, large, inconstant, ir- regular. Axial filament distinct in stained speci- mens, consisting of chitin or cutin-like substance. Flexible, elastic. Division transverse. Stains violet by Giemsa's stain and gray by iron-hemotoxylin. Cytoplasmic spirals stain with eosin, rubin, etc. Contain volutin granules. Trypsin digestion : Axial filament re- sistant. Bile salts (10 per cent) : Becomes shad- owy, pale, but is not dissolved. Saponin (10 per cent) : Lives 30 min- utes. Later becomes shadowy, but is not dissolved. Grows best under low oxygen tension. Optimum temperature 20° to 25°C. Habitat : Free-living in fresh or salt water. 2. Spirochaeta marina Zuelzer. (Spi- rochaeta plicatilis marina Zuelzer, Arch. f. Protistenk., ;24, 1912, 17; Zuelzer, ibid., 51 . ) From Latin, of the sea. Probably a subspecies or variant of Spirochaeta plicatilis. Cylindrical, 0.5 by 100 to 200 microns with blunt ends. Axial filament present. Flexible, elastic. Division transverse. Contains smaller and more irregularly distributed volutin granules than Spiro- chaeta plicatilis. Cj'toplasmic spirals stain. Grows best at low oxygen tension. Optimum temperature 20°C. Habitat : Sea water. 3. Spirochaeta eurystrepta Zuelzer. {Spirochaeta plicatilis eurystrepta Zuel- zer, Arch. f. Protistenk., 24, 1912, 17; FAMILY SPIROCHAETACEAE 1053 Zuelzer, ibid., 51 .) From Greek, broadly twisted . Probably a subspecies or variant of Spirochaeta plicatilis. Cylindrical : 0.5 micron in thickness and up to 300 microns in length. Spiral amplitude: More shallow than spirals of Spirochaeta plicatilis with blunt ends. Axial filament present. Fle.xible, elastic. Division transverse. Fewer volutin granules than in Spiro- chaeta plicatilis. Cytoplasmic spirals stain. Optimum temperature 20°C. Habitat : Swamp water and in grossly polluted water containing HjS. 4. Spirochaeta stenostrepta Zuelzer. (Zuelzer, Arch. f. Protistcnkunde, 24, 1912, 16; Treponema stenostrepta Ender- lein, Bakterien-Cyelogenie, 1925, 253.) From Greek, narrowly twisted. Cylindrical : 0.25 micron in thickness and 20 to 60, occasionally up to 200, microns in length, with pointed ends. Spiral amplitude very narrow with steep windings. Axial filament present. P'lexible, elastic. Division transverse. Fewer volutin granules than in Spiro- chaeta plicatilis. Cytoplasmic spirals stain. Optimum temperature 20"C. Habitat : Water containing H2S. 5. Spirochaeta daxensis Cantacuzene. (Compt. rend. Soc. Biol., Paris, 68, 1910, 75.) Named for Dax, a watering place in France. Large spirals: 0.5 to 2.5 by 30 to 100 microns, possessing a longitudinal chro- matin filament, and tapering at the ends. They are flattened and exhibit a double .series of curls, smaller waves being super- imposed on larger undulations. Optimum temperature 44° to 52°C. Source : Found in water of hot spring of Dax (52° to 56°C). Habitat : Hot springs. Appendix: The following species ma^' belong in this genus. Descriptions are usually inadequate. Spirochaeta agilis Adelmann. (Cent, f. Bakt., I Abt., Orig., 88, 1922, 413.) From mud from the harbor at Kiel. Original culture had a weak odor of hy- drogen sulfide. Spirochaeta aurantia Vinzent. (Compt. rend. Soc. Biol., Paris, 95, 1926, 1472.) From drain water. Forms small yellowish-orange colonies on agar after 5 to S days. Shows involution forms. This species definitely does not belong in the genus Spirochaeta, although it is placed here for the present . Its description sug- gests that it may belong among the vibrios. Spirochaeta flexihilis Xagler. (Cent, f. Bakt.. I Abt., Orig., 50. 1909, 445.) From the mud of a lake. Spirochaeta fulgurans Dobell. (Arch, f. Protistenk., 26, 1912, 117.) From water of the river Granta at Cambridge, ^vlay be a synonym of Spirochaeta steno- strepta. Spirochaeta gigantea Warming. (Warming, Videnskabl. Meddel. Xaturh. Foren., Copenhagen, 1875, 70; Spirillum giganteum Trevisan, I generi c le specie delle Batteriacee, 1889, 24; not Spirillum giganteum Migula, Syst. d. Bakt., i?, 1900, 1025.) Large, 3 microns in diameter. Inadequately described and possibly not a spirochaete. Spirochaeta graminea Zuelzer. (Zuel- zer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1670.) From an infusion of herbs. Size 0.25 by 100 microns. From rivers and marshes where it digests cellulose. Spirochaeta graminea marina Zuelzer. (Cent. f. Bakt., I Abt., Orig., 96, 1925, 426.) From sea water. Spirochaeta icterogenes marina Zuelzer. 1054 MANUAL OF DETERMINATIVE BACTERIOLOGY (Cent. f. Bakt., I Abt., Orig., 96, 1925, 426.) From sea water. Spirochaeta minima Dobell. (Dobell, Arch. f. Protistenk., 26, 1912, 117; not Spirochaeta viinima Pettit, Contribution a I'Etude des Spirochetides, Vanves, II, 1928, 187 {Treponema minimum Beaure- paire-Araguoand Vianna, Mem. Inst. Os- waldo Cruz, 5, 1913, 211).) One of the smallest known Spirochaeta, 0.5 by 2.0 to 2.5 microns. From water of the river Granta at Cambridge. Similar to Spiro- chaeta fulcjurans. Spirochaeta vivax (Dobell ) Zuelzer. {Treponema vivax Dobell, Arch. f. Pro- tistenk., 26, 1912, 117; Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., S, 1931, 1669.) From fresh water of the river Granta at Cambridge. Spirochaete kochii Trevisan. (Spiro- chaete des Wollsteiner See, Koch, in Cohn, Beilr. z. Biol. d. Pflanzen, 2, Heft 3, 1877, 420; Trevisan, Batter. Ital., 1879, 26; Spirillum kochii Trevisan, I generi e le specie delle Batteriacee, 1889, 24.) From water. Spirochaete schroeteri Cohn. (Jahres- ber. d. Schles. Gesellch. f. 1883, 198; quoted from Schroeter, in Cohn, Kryp- tog. Flora v. Schlesien, 3, 1, 1889, 168; Spirillum schroeteri Cohn, quoted from DeToni and Trevisan, in Saccardo, Syl- loge Fungorum, 8, 1889, 1007.) Similar to Spirochaete cohnii. From cellar walls. Genus II. Saprospira Gross. (Mittheil. Zool. Stat, zu Neapel, 30, 1911, 190.) Spiral protoplasm without evident axial filament. Spirals rather shallow. Trans- verse markings or septa (?) seen in unstained and stained specimens. Periplast membrane distinct. Motility active and rotating. Free-living in marine ooze. The type species is Saprospira grandis Gross. 1. Saprospira grandis Gross. (Mit- teil. Zool. Station zu Neapel, 20, 1911, 190.) From Latin, great. Cylindrical, 1.2 by SO microns in length, with obtuse ends. Spiral amplitude is 24 microns. Waves large, inconstant, shallow, ir- regular, 3 to 5 in number, sometimes al- most straight. Axial filament absent. Cross-striations i)rcsent. Membrane distinct. Division transverse. Flexible, elastic. Crista absent. Terminal spiral filament absent. Highly motile end portion absent. Trypsin digestion. Source : Found in intestinal tract of the oyster. Habitat : Free-living in foraminiferous sand. 2. Saprospira puncta Dimitroff. (Jour. Bact., 12, 1926, 146.) From Latin, pitted. Large spirals : 1 .0 by 86 microns with pointed ends. The spiral amplitude is 4 to 8 microns. The average number of turns is 3. Axial filament absent. Cross-striations present. Membrane distinct. Division transverse. Source : Found in oysters. 3. Saprospira lepta Dimitroff. (Jour. Bact., 12, 1926, 144.) From Latin, small. Large spirals : 0.5 by 70.0 microns, with pointed ends. The spiral amplitude ranges from 5 to 13 microns. The spiral width varies from 1.6 to 4.8 microns. The average number of turns is 6. Axial filament absent . Cross-striations present. FAMILY SPIROCHAETACEAE 1055 Membrane distinct. Division transverse. Source : Found in oysters in Baltimore, IMaryland . Appendix: Tlie following species have been placed in this genus. Saprospira flexuusa Dobell. (Arch. f. Protistenk., ;?6, 1912, 117.) Isolated once from water of the river Granta at Cam- bridge . Saprospira nana Gross. (IMittheil. Zool. Sta. zu Neapel, 20, 1911, 188.) From foraminiferous sand. Geniis III. Cristispira Gross. (Mittheil. Zool. Stat, zu Neapel, 20, 1910, 41.) Flexuous cell bodies in coarse spirals, 28 to 120 microns in length. Characterized by a crista or thin membrane of varying prominence on one side of the body extend- ing the entire length of the organism. Cross-striations. Actively motile. Found in the intestinal tract of molluscs. The type species is Cristispira balbianii (Certes) Gross. 1. Cristispira balbianii (Certes) Gross. {Trypanosoma balbianii Certes, Bull. Soc. Zool. de France, 7, 1882, 347; Spiro- chaeta balbianii Swellengrebel, Ann. Inst. Past., 21, 1907, 562; Spirochaete balbianii Borrel and Cernovodeanu, Compt. rend. Soc. Biol., Paris,' 6'^, 1907, 1102; Gross, Cent, f . Bakt., I Abt., Grig., 65, 1912, 90.) Cylindrical: 1.0 to 3.0 by 40 to 120 microns, with obtuse ends. Spiral amplitude is 8 microns. Spiral depth is 1.6 microns. Waves 2 to 5, sometimes more, large, irregular, shallow. Axial filament absent . Cross-striations present. JNIembrane distinct. Flexible, elastic. Crista present, a ridge-like membrane making one to two complete turns. Terminal spiral filament absent. Highly motile end portion absent. Stains : Cell membrane behaves like chitin or cutin substance. Stains violet by Giemsa's solution, and light graj- by iron-hemotoxylin. Trypsin digestion: Membrane resist- ant, crista and striations disappear. Bile salt (10 per cent) : Crista quickly dissolves. Saponin (10 per cent) : Crista becomes fibrillar, then indistinct. Source : From the crystalline style of oysters. Habitat : Parasitic in alimentary tract of shell-fish. 2. Cristispira anodontae (Keysselitz) Gross. (Spirochaeta anodontae Keysse- litz, Arb. a. d. kaiserl. Gesundheitsamte, 23. 1906, 566 ; Gross, Cent . f. Bakt., I Abt., Grig., 65, 1912, 900.) From M.L., mus- sels. 0.8 to 1.2 by 44 to 88 microns with sharply pointed ends; flattened and possessing an undulating membrane. The periplast is fibrillar in appearance and there is a dark granule at each end of the undulating membrane. The chro- matin material is distributed in the form of globules or elongated bands. Large spirals : The average width of the spiral is 2 microns. The average wave length is 8 microns. The number of complete turns ranges from 5 to 1 1 . Habitat : Found in the crystalline style of fresh water mussels, Anodonta cygnea and .4. tnutabilis, also in intestinal tract of oysters. 3. Cristispira pinnae (Gonder) Bergey et al. {Spirochaete pinnae Gonder, Cent, f. Bakt., I Abt., Grig., 47, 1908, 491; Spirochaeta pinnae Schellack, Arb. a. d. 1056 MANUAL OF DETERMINATIVE BACTERIOLOGY kaiserl. Gesundheitsamte, 30, 1909, 379; Bergey etal., Manual, 1st ed., 1923, 423.) From Latin, of a mussel. Spirals: 0.5 to 3.0 by 10 to 60 microns, round in section with blunt ends, the one being slightly more pointed than the other. They have a ridge or comb running along one side but no terminal filaments. Cross-striations distinct. The chromatin grarmles are grouped in fours. An undulating membrane can be dem- onstrated. Source : Found in the intestinal canal of the scallop (Pecten jacobaeus) . Habitat: From the crystalline style of molluscs. Appendix: Additional species which ap- pear to belong in this genus are : Cristispira acuminata (Schellack) Ford. {Spirochaeta acuminata Schel- lack, Arb. kais. Gesundheitsamte, 30, 1909, 379; not Spirochaeta acuminata Castellani, Brit. Med. Jour., 2, 1905, 1330; Ford, Textb. of Bact., 1927, 939.) From the crystalline style of a mollusc. Tapes laeta. Cristispira cardii-papillosi (Schellack) Ford. {Spirochaeta cardii-papillosi Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Ford, Textb. of Bact., 1927, 939.) From the crystalline style of a mollusc, Cardium papillosum. Cristispira chamae (Schellack) Nogu- chi. {Spirochaeta chamae Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Noguchi, Jour. Exp. Med., 27, 1918, 583.) From the crystalline styles of molluscs, Chama spp. Cristispira gastrochaenae (Schellack) Ford. {Spirochaeta gastrochaenae Schel- lack, Arb. kais. Gesundheitsamte, SO, 1909, 379; Ford, Textb. of Bact., 1927, 940.) From a shellfish, Gastrochaena dubia. Constant length 29 microns. Cristispira helgolandica Collier. (Cent. f. Bakt., I Abt., Orig., 86, 1921, 132.) Found three times in the body fluid of an echinoderm, Asterias rubens, in the North sea. Average length 68 microns. Named for the place where the investigation was made (Helgoland). Cristispira interrogationis Gross. (Mit- theil. Zool. Station zu Neapel, 20, 1910, 41.) From the intestinal canal of the scallop, Pecten jacobaeus. Cristispira limae (Schellack) Ford. {Spirochaeta limae Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Ford, Textb. of Bact., 1927, 939.) From the crystalline styles of molluscs, JAma spp. Similar to Cristispira balbianii. Cristispira mactrae (Prowazek) Ford. {Spirochaeta mactrae Prowazek, Arch. f. Schiffs- u. Tropenhyg., 14, 1910, 297; Ford, Textb. of Bact., 1927, 940.) From the digestive tract of a shellfish, Mactra sulcataria. Cristispira mina Dimitrofi'. (Jour. Bact., 12, 1926, 159.) Found in oysters. Cristispira modiolae (Schellack) Nogu- chi. {Spirochaeta modiolae Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379 ; Noguchi, Jour. Exp. Med., 27, 1918, 583.) Found in mussels and oysters. Cristispira ostrcae (Schellack) Nogu- chi. {Spirochaeta ostreae Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Noguchi, Jour. Exp. Med., 27, 1918, 583.) From the crystalline style of the oyster, Ostrea edulis. Identical with Cristispira anodontae Gross. Cristispira pachelabrae de Mello. (Compt. rend. Soc. Biol., Paris, 84, 1921, 241 .) From the digestive tract of a shell- fish, Pachelahra moestra. Cristispira parvula Dobell. (Arch. f. Protistenk., 26, 1912, 117.) From the crystalline style of a mollusc, Venus {Meretrix) castra, in Ceylon. The small- est Cristispira known — 0.4 to 0.5 by 20 to 45 microns. Cristispira pectinis Gross. (Mittheil. Zool. Sta. zu Neapel, .20, 1910, 41.) From the digestive tract of a scallop, Pecten jacobaeus. Identical with Cristispira balbianii Gross. Cristispira polydorae Mesnil and Caul- FAMILY SPIROCHAETACEAE 1057 lery. (Compt. rend. Soc. Biol., Paris, 75, 1916, 1118; Cristispirella polydorae Hollande, Compt. rend. Acad. Sci. Paris, 173, 1921, 1696). From a marine annelid, Polydora flava. Cristispira pusilla (Schellack) Ford. (Spirochaeta pusilla Schellack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Ford, Textb. of Bact., 1927, 940.) From the digestive tract of a mussel, Anodnnfa mutabilis. Cristispira saxicavae (Schellack) P^ord. {Spirochaeta saxicavae Schellack, Arb. kais. Gesundheitsamte, SO, 1909, 379; Ford, Textb. of Bact., 1927, 940.) From the crystalline style of a mollusc, Saxi- cava arctica. Cristispira spiculifera (Schellack) Di- mitroff. (Spirochaeta spiculifera Schel- lack, Arb. kais. Gesundheitsamte, 30, 1909, 379; Dimitroff, Jour. Bact., 12, 1926, 157.) Found in mussels. Cristispira tapetos (Schellack) Gross. {Spirochaeta tapetos Schellack, Arb. kais. Gesundheitsai te, 30, 1909, 379; Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 84.) From the crystalline style of a mollusc. Tapes decussata. Cristispira tenua Dimitroff. (Jour. Bact., 13, 1926, 160.) Found in oysters. Cristispira veneris Dobell. (Quart. Jour. Microsc. Sci., London, 54, 1910- 1911, 507 and ibid., 56, 1911, Part 3.) From a clam, Veiius {Meretrix) castra, in Ceylon. Identical with Cristispira bal- hianii Gross. Cristispirella caviae Hollande. (Compt. rend. Acad. Sci., Paris, 172, 1921, 1693.) From the intestine of a guinea pig. Probably a protozoan. Evi- dently the same as Heliconema (see ap- pendix to Borrelia). Both Cristispirilla polydorae and Cristispirella caviae have characteristics at variance with accepted ideas of spirochaetes. Spirochaeta solenis Fantham. (Ann. Trop. Med. and Parasitol., 5, 1911, 479.) A parasite of a mollusc, Solen ensis. 1058 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY II. TREPONEMATACEAE SCHAUDINN. (Deutsche med. Wochnschr., 31, 1905, 1728.) Coarse or slender spirals, 4 to 16 microns in length; longer forms due to incomplete or delayed division. Protoplasm with no obvious structural features. Some may- show terminal filaments. Spirals regular or irregular, flexible or comparatively rigid. Some visible only with dark field illumination. Parasitic on vertebrates with few exceptions. Some pathogenic. Many can be cultivated. Key to the genera of family Treponemataceae. I. Stains easily with ordinary aniline dyes. Genus I. Borrelia, p. 1058. II. Stain with difficulty except with Giemsa's stain and silver impregnation. A. Strict anaerobes. Genus II. Treponema, p. 1071. B. Aerobes. Genus III. Leptospira, p. 1076. Genus I. Borrelia* Sivellengrebcl . (Swellengrebel, Ann. Inst. Past., £1, 1907, 582; Spiroschaudinnia Sambon, in Man- son, Tropical Diseases, August, 1907, 833; Cacospira Enderlein, Sitzber. Ges. Naturf. Freunde, Berlin, 1917, 309; Entomospira Enderlein, ibid.; Spironema Bergey et al., Manual, 1st ed., 1923, 424; not Spironema Vuillemin, Compt. rend. Acad. Sci. Paris, 140, 1905, 1567; Spirochaeta Gieszczykiewicz, Bull. Acad. Polonaise d. Sci. et Lettres, CI. Sci. Math, et Nat., Ser. B, 1939, 24.) Length 8 to 16 microns. Coarse, shallow, irregular, with a few obtuse angled spirals. Generally taper terminally into fine filaments. Stain easily with ordinary aniline dyes. Refractive index approximately the same as that of true bacteria. Parasitic upon many forms of animal life. Some are pathogenic for man, other mammals and birds. Generally hematophytes are found on mucous membranes. Some are trans- mitted by the bites of arthropods. The type species is Borrelia anserina (Sakharofi) Bergey et al. 1. Borrelia anserina (Sakharofi) Ber- SjMionyms: SpirocJtueta marchotixi gey et al. (Spirochaeta anserina Sak- Xuttall, Epidemiol. Soc, London, £4, haroff, Ann. Inst. Past., 5, 1891, 564; 1904, 12 (Spirille de la poule, Marchoux Spirillum anserum (sic) Sternberg, Man. and Salimbeni, Ann. Inst. Past., 17, 1903, of Bact., 1893, 499; Spirillum anserinum 569; Spirochaeta gallinarum Stephens Mac^, Traite Pratique de Bact., 4th ed., and Christopher, Practical Study of Ma- 1901, 1060; Spirochaete anserina Mace, laria and Other Blood Parasites, Liver- ihid.; Spiroschaudinnia anserina Cas- pool, 1905; Borrelia gallinarum Swellen- tellani and Chalmers, Man. Trop. Med., grebel, Ann. Inst. Past., 21, 1907, 623; 2nd ed., 1913, 403; Spironema anserina Spirochaete gallinarum Lehmann and Noguchi, Jour. Exp. Med., 27, 1918, 584; Neumann, Bakt. Diag., 4 Aufl., 2, 1907, Bergey et al.. Manual, 2nd ed., 1925, 435 ; 623 ; Spironema gallinarum Gross, Cent. f. Treponema anserina Noguchi, in Jordan Bakt., I Abt., Grig., 65, 1912, 92; Spiro- and Falk, Newer Knowledge Bact. and schaudinnia marchouxi Castellani and Immun., 1928, 456.) From Latin, per- Chalmers, Man. Trop. Med., 2nd ed., taining to geese. 1913, 403; Spironema marchouxi Ford, * Further revision of the genus by Prof. E. G. D. Murray, McGillUniv., Montreal, P. Q., Canada, April, 1947. Reviewed by Dr. Gordon E. Davis, Rocky Mountain Laboratory, U.S.P.H.S, Hamilton, Montana. FAMILY TREPONEMATACEAE 1059 Textb. of Bact., 1927, 955; Spirochaeta gallinae Ford, idem; Treponema gallin- arum Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 461 ; Treponema marchouxi Gay et al., Agents of Disease and Host Resistance, 1935, 1077). The cause of septicaemia in chickens. Spirochaeta granulosa perietrans Bal- four, Jour. Trop. Med. and Hyg., 10, 1907, 153 {Spiroschaudinnia granulosa Balfour, Jour. Trop. Veter., Calcutta, 5, 1910, 309 ; Spironema granulosa Ford, Textb. of Bact., 1927, 957). From spirochetosis of fowls in Sudan. Spirochaeta nicollei Brumpt, Bull. Soc. Path. Exot., 2, 1909, 285 and/or Precis de Parasitol., Paris, 1st ed., 1910 (Galli- Valerio, Cent. f. Bakt., I Abt., Orig., 50, 1909, 189 and 61, 1912, 529; Spir a neyna nicollei Ford, Textb. of Bact., 1927, 958; Treponema nicollei Gay et al., Agents of disease and Host Resistance, 1935, 1077). From spirochetosis of geese in Tunisia. Spirochaeta neveuxi Brumpt, Bull. Soc. Path. Exot., 2, 1909, 285 {Spiroschaudin- nia neveuxii Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404; Spironema neveuxi Ford, Textb. of Bact., 1927, 958; Treponema neveuxi Gay et al.. Agents of Disease and Host Resistance, 1935, 1077). The cause of fowl spiroche- tosis in Senegal. Spirochaeta gallinarum var. hereditaria Neumann and IMayer, in Lehmann, Med. Atlanten, 11, 1914, 276. A North African strain of fowl spiz'ochetosis. Borrelia pullorum Redowitz, Amer. Jour. Med. Technol., 2, 1936, 91. From diseased chickens. Spirochaeta anatis Parrot, Bull. Soc. Path. Exot., 13, 1920, 647. Pathogenic for domestic ducks in Algeria. Morphology: 0.25 to 0.3 by 8 to 20 microns, averaging about 1 spiral per micron. Actively motile, with lashing move- ments. Stains readily with aniline dyes and Giemsa's stain. Cultivation : Can be cultivated in Noguchi's ascitic fluid-rabbit kidney medium. Immunologj' : Antigenically distinct from species found in mammals. Arthropod vectors : Transmitted by the bites of ticks (Argas persicus, A. minia- tus, A. reflexus and Ornithodoros mou- hata.) Pathogenic for birds but not for mam- mals. Source: From blood of infected geese, ducks, fowls and vector ticks. Habitat : The cause of spirochetosis of fowls. 2. Borrelia recurrentis (Lebert)Bergey et al. (Obermeier, Berlin, klin. Woch- schr., 1873, 152; Protomycetum recurrentis Lebert, Ziemssen's Handbuch, 2, 1874, 267; Spirochaete obermeieri Cohn, Beitr. z. Biol. d. Pdanzen, 1, Heft 3, 1875, 196; Spirillum obermeieri Zopf, Die Spalt- pilze, 3 Aufl., 1885, 71 ; Spirochaeta ober- meieri Migula, in Engler and Prantl, Die natiirl. Pflanzenfam., 1, la, 1895, 35; Spirochaete recurrentis Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 621 ; Spirochaeta recurrentis Castellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 305; Spironema recurrentis Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 85; Spiroschaudinnia recurrentis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 398; Spironema obermeieri Park and Williams, Pathogenic Microorganisms, 6th ed., 1917, 513; Cacospira recurrentis Enderlein, Sitzungsber. d. Gesellsch. naturf. Freunde, 1917, 313; Treponemare- currentis Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 508; Treponema obermeieri Brumpt, ibid.; Cacospira ober- meieri Enderlein, Bakterien-Cyclogenie, 1925, 254; Bergey et al.. Manual, 2nd ed., 1925, 433; Spirillum recurrentis Ford, Textb. of Bact., 1927, 9i8; Spiroschaudin- nia obermeieri Ford, ibid.) From Latin, recurring. Cylindrical or slightly flattened, 0.35 to 0.5 by 8 to 16 microns, with pointed ends. 1060 MANUAL OF DETERMINATIVE BACTERIOLOGY Spiral amplitude 1.5 microns. Spirals large, wavy, inconstant, about 5 in number. Terminal finely spiral filaments present. Highly motile end portion absent. Motility : By active cork-screw motion without polarity. Lashing movements common in drawn blood. Stains with common aniline dyes. Gram-negative. Violet with Giemsa's stain. Bile salts (10 per cent) : Disintegration complete. Saponin (10 per cent) : Immobilized in 30 minutes, then broken up in a few hours. In some a skeletal structure remains. Cultivation : Can be cultured in ascitic or hydrocoel fluid to which a piece of sterile rabbit kidney is added . Optimum reaction pH 7.2 to 7.4. Immunology : Serum does not agglu- tinate Borrelia duttoni. Accidental and experimental transmis- sion by conjunctival sac and skin abra- sions. Disease in experimental animals (small rodents after monkey passage) mild. Arthropod vector: Louse {Pediculus humanus) which exhibits normal trans- mission from the 16th to the 28th day. Found in the bed-bug {Cimex lectularius) and ticks, but not transmitted by them. No evidence of hereditary transmission in the louse. Habitat: The cause of European re- lapsing fever. Transmissible to man, monkeys, mice and rats. 3. Borrelia duttonii (Breinl) Bergey et al. (Dutton and Todd, Brit. Med. Jour., 2, 1905, 1259; Spirochaeta duttoni Breinl, Lancet, 2, 1906, 1690; Spirillum duttoni Novy and Knapp, Jour. Inf. Dis., 3, 1906, 296; see Dutton and Todd, Jour. Trop. Med., 10, 1907, 385; Spirochaete duttoni Lehmann and Neumann, Bakt. Diag., 4 Aufl., 2, 1907, 623; Spironema duttoni Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 94; Spirochaeta Micro- spironema duttoni Duboscq and Lebailly, Compt. rend. Acad. Sci., 154, 1912, 662; Spiroscliaudinnia duttoni Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 399; Treponema duttoni Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922, 497; Cacospira duttoni Enderlein, Bakterien-Cyclogenie, 1925, 254; Bergey et al.. Manual, 2nd ed., 1925, 434.) Named for Dutton, who discovered this organism. Morphology : Similar to Borrelia recur- rentis. Cultivation : Growth occurs under anaerobic conditions in serum water, hydrocoel or ascitic fluid to which a piece of sterile rabbit kidney is added. Immunology : This organism is anti- genically distinct from other causes of relapsing fever. Pathogenic for mice and rats. Disease in small rodents and many other experi- mental animals very severe. Arthropod vector: This species is transmitted to man through the bite of the tick {Ornithodoros moubata) by fecal contamination of the bite. In the tick the organism goes through some granula- tion or fragmentation phenomenon, the nature of which is not understood. Hereditary transmission to at least the third generation of the tick. Not trans- mitted by the louse. Habitat: The cause of Central and South African relapsing fever. 4. Borrelia kochii (Novy) Bergey etal. {Spirochaeta kochi Novy, Proc. Path. Soc. Philadel., N. S. 10, 1907, 1; Spirochaeta rossi Nuttall, Jour. Roy. Inst. Pub. Health, London, 16, 1908, 385; Spiro- scliaudinnia rossii Castellani and Chal- mers, Man. Trop. Med., 2nd ed., 1913, 400; Spironema kochii Noguchi, Jour. Exp. Med., 27, 1918, 584; Treponema kochi Brumpt, Nouveau Traitd de M6de- cine, Paris, 4, 1922, 497; Treponema rossi Brumpt, ibid.; Bergey et al., Manual, 2nd ed., 1925, 434; Borrelia rossi Stein- haus, Insect Microbiology, 1946, 452.) FAMILY TREPONEMATACEAE 1061 Named for Koch, who first observed spirochetes in East African relapsing fever. Morphology : Similar to that of Borrelia recurrentis. Cultivation : Same as for Borrelia re- currentis. Immunology : Antigenically distinct from both Borrelia recurrentis and B. duttonii. Pathogenic for mice and rats. Arthropod vector: No record. Habitat : The cause of African relapsing fever. 5. Borrelia novyi (Schellack) Bergey et al. (Spirochaete from relapsing fever, Norris, Pappenheimer and Flournoy, Jour. Inf. Dis., 3, 1906, 266; Spirochaeta novyi Schellack, Arb. kaiserl. Gesund- heitsamte, £7, 1907, 199 and 364; Spiro- nema novyi Gross, Archiv f. Protistenk., £4, 1912, 115; Spire schaudinnia novyi Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 400; Treponema novyi Brumpt, Nouveau Traits de M4de- cine, Paris, 4, 1922, 508; Cacospira novyi Enderlein, Bakterien-Cyclogenie, 1925, 254; Bergey et al., JNIanual, 2nd ed., 1925, 434.) Named for Novy, the American bacteriologist. Morphology : Similar to that of Borrelia recurrentis. Cultivation: Same as for Borrelia re- currentis. Immunology : Antigenically distinct from other relapsing fever organisms. Pathogenic for monkeys, white rats and white mice. Arthropod vector: Unknown. Habitat: Recovered from a patient in Belle vue Hospital, New York. Origin of infection unknown. 6. Borrelia berbera (Sergent and Fo- ley) Bergey et al. {Spirochaeta berbera Sergent and Foley, Ann. Inst. Past., 24, 1910, 337; Spiroschaudinnia berbera Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 402; Spironema berbera Noguchi, Jour. E.xp. Med., 27, 1918, 584; Spirochaeta berbera Kolle and Hetsch, Exper. Bakt. u. Infekt., 6 Aufl., 1, 1922, 811; Treponema berberum Brumpt, Nou- veau Traits de M^decine, Paris, 4, 1922, 496; Bergey et al.. Manual, 2nd ed., 1925, 435.) Named for the Berbers, a tribe of Northern Africa. Morphology : More tenuous than other relapsing fever organisms, 0.2 to 0.3 by 12 to 24 microns. Cultivation: No record of its cultiva- tion. Immunology : Antigenically distinct from Borrelia recurrentis. Arthropod vector : Possibly carried by the louse (Pediculus vestimenti). Source : Found in cases of relapsing fever in Algiers, Tunis and Tripoli. Habitat : Cause of relapsing fever in North Africa. Is virulent for monkeys. Produces non-fatal infections in rats and mice. 7. Borrelia carteri (Mackie) Bergey et al. (Spirochaeta carteri Mackie, Ann. Trop. Med. and Parasitol., 1, 1907, 157 and Indian Med. Gazette, 44, 1908, 370; Spirillum carteri Mackie, Lancet, 2, 1907, 832, according to Ford, Textb. of Bact., 1927, 950; Spiroschaudinnia carteri Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 401; Spironema carteri Noguchi, Jour. Exp. Med., 27, 1918, 584; Treponema carteri Brumpt, Nouveau Traite de M^decine, Paris, 4, 1922, 497; Bergey et al.. Manual, 2nd ed., 1925, 435.) Named for Carter, who in 1879 described this organism in the blood of patients with Indian relapsing fever. Morpholog}" : Similar to Borrelia ber- bera. Cultivation: Not recorded. Immunology : Probably a distinct spe- cies. A succession of distinct .serological types occurs with the relapses in a single infection (Cunrdngham et al., Far East- ern Association of Tropical Medicine, Tokj'o, ;^1925; Indian Journal of Medical 1062 MANUAL OF DETERMINATIVE BACTERIOLOGY Research, 22, 1934-1935, 105 and 595; iUd., 24, 1937, 571 and 581). Arthropod vector: Carried by either Pediculus vestimenti ov Cimex rolundatus or by both. Habitat: The cause of Indian relapsing fever. Transmissible to monkeys, rab- bits, rats and mice. 8. Borrelia theileri (Laveran) Bergey et al. {Spirochaela theileri Laveran, Compt. rend. Acad. Sci., Paris, 136, 1903, 939; Spiroschaudinnia theileri Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404; Spironema theileri Noguchi, Jour. Exp. Med., 27, 1918, 584; Bergey et al., Manual, 2nd ed., 1925, 435; Spirillum theileri and Spirochaete theileri Pettit, Contribution a I'Etude des Spiro- ch^tidds, Vanves, II, 1928; Treponema theileri Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 461.) Named for Theiler, who discovered this organism in 1902 in Transvaal, South Africa. Morphology : 0.25 to 0.3 micron by 20 to 30 microns with pointed ends. Cultivation: No record. Immunology : Is distinct from the spe- cies infecting man. Arthropod vector: Transmitted by the tick {Rhipicephalus decoloratus) . Source : Blood of cattle. Habitat : Blood of cattle and other mammals in South Africa. 9. Borrelia glossinae (Novy and Knapp) Bergey et al. {Spirillum glossi- nae Novy and Knapp, Jour. Inf. Dis., 8, 1906, 385; Spirochaeta glossinae Cas- tellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 310; Spiroschaudinnia glossinae Castellani and Chalmers, ibid., 3rd ed., 1919, 454; Spironema glossinae Bergey et al.. Manual, 1st ed., 1923, 425; Bergey et al., Manual, 2nd ed., 1925, 435; Entomospira glossinae Enderlein, Bak- terien-Cyclogcnie, 1925, 254; Treponema glossinae Ford, Te.xtb. of Bact., 1927, 988.) Named for the genus of insects, Glossina. Morphology : 0.2 \)y 8.0 microns, oc- curring singly, sometimes in pairs. Gen- erally 4 spirals. Shorter, narrower and has more turns than has Borrelia rec2ir- rentis. Habitat : Found in the stomach con- tents of the tse-tse fly {Glossina pal- palis ) . 10. Borrelia buccale (Steinberg) Brumpt. (Spirochaeta buccalis Stein- berg, 1862, according to Hoffmann and Provvazek, Cent. f. Bakt., I Abt., Orig., 41, 1906, 819; Spirochaete cohnii Winter, Die Pilze, 1879, 61; (?) Microspira buc- calis Lewis, The Lancet, 1884, quoted from Schroeter, in Cohn, Kryptog. Flora V. Schlesien, 3, 1, 1889, 169; Spirochaete buccalis, quoted from Schroeter, ibid., 168; Spirillum cohnii Trevisan, I generi e le specie delle Batteriacce, 1889, 24; Spirillum buccale Mac6, Traite Pratique de Bact., 4th ed., 1901, 1062; Spirochaeta inaequalis Gerber, Cent. f. Bakt., I Abt., Orig., 56, 1910, 508; Spirochaeta undu- lata Gerber, idem; Treponema buccale Dobell, Arch. f. Protistenk., 26, 1912, 117; Spironema buccale Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 84; Spiro- schaudinnia buccalis Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 450; Brumpt, Nouveau Trait(5 de M^de- cine, Paris, 4, 1922, 495; Treponema in- aequale Brumpt, ibid.; Treponema un- dulatum Brumpt, ibid., 514.) From Latin buccalis, buccal. Morphology: 0.4 to 0.9 by 7 to 20 mi- crons. The largest of the mouth spiro- chetes. Motility: Active, serpentine, rotating and fiexuous. Staining: Stains with aniline dyes and is violet with Giemsa's stain. Cultivation : Has not been obtained in pure culture and probably does not grow in any medium tried to date. Habitat: In normal mouths and invades formed lesions of the respiratory mucous membrane. FAMILY TREPONEMATACEAE 1063 11. Borrelia vincentii (Blanchurd) Ber- gej' ot al. (Spirorhacta vincenli Blan- chard, Arch. f. Protistenk., 10, 1906, 129; Spirochaeta schaudinni Prowazek, Arb. kaiserl. Gesundheitsamte, 22, 1907, 23; Spirochaete plaut-vincenti Lehmann and Neumann, Bakt. Diag., 5 Aufl., 2, 1912, 579; Spiroschaudinnia vincenli Castel- lani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 402; Spiroschaudinnia schaudinni Castellani and Chalmers, idem; Spironema vincenli Park and Williams, Pathogenic Microorganisms, 6th ed., 1917, 506; Treponema vincenli Brumpt, Nouveau Traits de M^decine, Paris, J^, 1922, 514; Treponema schaudinni Brumpt, idem; Bergey et al.. Manual, 2nd ed., 1925, 435.) Named for Vincent, the French bacteriologist. Morphology: 0.3 by 8 to 12 microns, 3 to 8 irregular shallow spirals. Stains easily with the common aniline dyes and is Gram -negative. Motility : Has a rapid progressive and vibratory motion. Cultivation : Can be cultivated under anaerobic conditions. Cultures may show long forms with only a writhing motion. Not pathogenic for laboratory animals. Habitat : Found on normal respiratory mucous membrane and is associated with a fusiform bacillus {Fusobaclerium plauli-vincenti) in Vincent's angina. 12. Borrelia refringens (Schaudinn and Hoffmann) Bergey et al. (Spiro- chaeta refringens Schaudinn and Hoff- mann, Arbeiten kaiserl. Gesundheits- amte, 22, 1905, 528; Spirochaeta refrin- gens Hoffmann and Prowazek, Cent. f. Bakt., I Abt., Orig., 41, 1906. 7-12; Spiro- nema refringens Gross, Arch. f. Protis- tenk., 24, 1912, 115; Spiroschaudinnia refringens Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 459; Trepo- nema refringens Castellani and Chalmers, ibid., 461; Bergey et al.. Manual, 2nd ed., 1925, 436.) From Latin, refractive. Morphology: 0.5 to 0.75 by 6 to 20 mi- crons. Spirals are coarse and shallow. Spirals are generally smoothly lounded and regular, tapering towards the end into a fine projection. Stains easily by common dyes. In stained specimens the spirals appear irregular. Motility: Active serpentine and rotat- ing motion with marked flexion. Cultivation: Uncertain. Pathogenicity: None. Source : Found with Treponema palli- dum in some cases of syphilis as orig- inally described by Schaudinn. Habitat : Genital mucous membranes and necrotic lesions of the genitalia of man . 13. Borrelia hyos (King and Drake) Bergey et al. (Hog cholera virus, King and Baeslack, Jour. Inf. Dis., 12, 1913, 39; Spirochaeta suis King, Baeslack and Hoffmann, Jour. Inf. Dis., 12, 1913, 2.35; not Spirochaeta suis Bosanquet, Spiro- chetes, Saunders, 1911; Spirochaeta hyos King and Drake, Jour. Inf. Dis., 16, 1915, 54 ; Spironema hyos Bergey et al., Manual, 1st ed., 1923, 426; Bergey et al., Manual, 2nd ed., 1925, 436; Spironema suis Ford, Textb. of Bact., 1927, 959.) From Greek, hog. Morphology : 1 micron bj' 5 to 7 microns. Distinctly shorter and thicker than other members of the genus. Motility: .\ctive spinning motion, spirals fixed. Cultivation : Grows under anaerobic conditions in the presence of tissue. Habitat : Found in the blood, intestinal ulcers and other lesions of hogs suffering from hog cholera. 14. Borrelia hermsi (Davis) Steinhaus. (Spirochaeta hermsi Davis, Amer. Assoc. Adv. Sci., Pub. No. 18, 1942, 46; Stein- haus, Insect Microbiology, 1946, 453.) Investigations by Davis (loc. cit.) indi- cate that each species of Ornilhodoros that is a relapsing fever vector carries a spirochete that is tick-host specific and that this host-specific relationship offers a more accurate approach to the differen- tiation of relapsing fever spirochetes 1064 MANUAL OF DETERMINATIVE BACTERIOLOGY than any of the several criteria pre- viously used. This was shown to be the case for Borrelia hermsi and Borrelia parkeri. For this reason no attempt is made to describe the morphology and other char- acters of the relapsing fever spirochetes of North and South America. Borrelia hermsi is transmitted by Ornithodoros hermsi. A cause of relapsing fever in the West- ern part of the U. S. A. 15. Borrelia parkeri (Davis) Steinhaus. (Spirochaeta parkeri Davis, loc. cii.; Steinhaus, loc. cit.) Transmitted by Ornithnodoros parkeri. A cause of relapsing fever in the West- ern part of the U. S. A. 16. Borrelia turicatae (Brumpt) Stein- haus. (Spirochaela luricalae Brumpt, Comp. rend. Soc. Biol., Paris, llS, 1933, 1369; Steinhaus, loc. cit.) Transmitted by Ornithodoros turicala. A cause of relapsing fever in Mexico, Texas and nearby areas. 17. Borrelia venezuelensis Brumpt. {Treponema venezuelensis Brumpt, Nou- veau Traite de M(5decine, Paris, 4, 1922, 492; Brumpt, ibid., 495; Spirochaela ven- ezuelensis Pettit, Contributions al'Etude des Spiroch(5tides, Vanves, ^,1928, 295.) Transmitted by Ornithodoros rudis (0. venezuelensis). A cause of South American relapsing fever. Brumpt (Precis dc Parasitologic, 3rd ed., Paris, 1936) regards this species as identical with Borrelia neotropicalis (Bates, Dunn and St. John) Steinhaus. {Treponema neotropicalis Bates, Dunn and St. John, Amer. Jour. Trop. Med., 1, 1921, 183; Spirochaeta neotropicalis St. John and Bates, Amer. Jour. Trop. Med., 2, 1922, 251; Steinhaus, loc. cit.) Transmitted by Ornithodoros venezuelen- sis. A cause of relapsing fever in Panama. Appendix: Many of the species in- cluded in this appendix are so inade- quately described that it is not certain that they belong in this group. Borrelia phagedenis (Noguchi) Bergey et al. {Spirochaeta phagedenis Noguchi, Jour. Exp. Med., 16, 1912, 2Ql;'Spiro- schaudinnia phagedenis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 403 ; Treponema phagedenis Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922, 511 ; Spironema phagedenis Bergey et al.. Manual, 1st ed., 1923, 426; Bergey et al.. Manual, 2nd ed., 1925, 435.) From phagedenous ulcer. Heliconema pyrphoron Scholer. (Cent. f. Bakt., I Abt., Orig., 138, 1937, 342.) From human blood. Pathogenic. Heliconema vincenti Sanarelli. (Ann. Inst. Past., 41, 1927, 701.) From the in- testine of a guinea pig. Shows stages be- tween spirochetes and fusiform bacilli. (See Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 130.) Microspironema merlangi Duboscq and Lebailly. (Compt. rend. Acad. Sci. Paris, 154, 1912, 662.) From the whiting, Merlangus merlangus. May bo a syn- onym of Spirochaeta gadi. Spirillum, gondii Nicolle. (Nicollo, Compt. rend. Soc. Biol., Paris, 63, 1907, 213; Spirochaeta gondi Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 16S0.) Found in the blood of a rodent, Ctenodactylus gondi. Not patho- genic. Associated with a piroplasma. Probably not a spirochete. Spirillum lalapiei Laveran. (Laveran, Bull. Soc. Path. Exot., 1, 190S, 148; Spirochaela lalapiei Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1683 ; Spironema latapici (sic) Ford, Textb. of Bact., 1927, 964.) From the blood of a shark. Spirillum pithed Thiroux and Du- fouger^. (Thiroux and Dufourger(5, Compt. rend. Acad. Sci. Paris, 150, 1910, 132; Spirochaela pitheci Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1676; Spironema pitheci Ford, FAMILY TREPONEMATACEAE 1065 Textb. of Bact., 1927, 961.) From the blood of an African monkey, Ccrcopithe- cus patas. Pathogenic for monkeys, rats and field mice. Closely related to Boi-- relia duttonii. Spirochaeta aboriginalis Cleland. (Cleland, Jour. Trop. Med., 12, 1909, 143; Spiroschaudinnia aboriginalis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 402; Treponema aboriginalis Brumpt, Xouveau Traite de Medeeine, Paris, ^, 1922, 496.) Found in cases of granuloma inguinale in West Australia. Probably saprophytic. Spirochaeta acuminata Castellani. (Castellani, Brit. Med. Jour., 2, 1905, 1330; Spirochaeta tenuis acuminata Cas- tellani, idem and/or Arch. f. Schiffs- u. Tropenhj'g., 12, 190S, 311 ; Spiroschaudin- nia acuminata Castellani and Chalmers, Man. Trop. Med., 3rd ed;, 1919, 449; Treponema acuminatum Brumpt, Nou- veau Traits de Medeeine, Paris, 4, 1922, 496.) From ulcerated lesions of yaws. Spirochaeta acuta Kritchewski and Seguin. (Rev. de StomatoL, 22, 1920, 613.) From the oral cavity. Spirochaeta aeglejlni Henry. (Jour. Path, and Bact., IS, 1913, 222.) From haddock. Spirochaeta aegyptica Gonder. (Gon- der, in Prowazek, Handb. d. path. Pro- toz., 6, 1914, 671; Spironema aegyptica Noguchi, Jour. E.xp. Med., 27, 1918, 584; Treponema egypticum Brumpt, Nouveau Traits de Medeeine, Paris, 4, 1922, 500; Borrelia aegypticum Steinhaus, Insect Microbiology, 1946, 452.) Observed in cases of relapsing fever in Sudan. Prob- ably a S3'nonym of Borrelia recurrentis . Spirochaeta ambigua Seguin and Vin- zent. (S(5guin and Vinzent, Compt. rend. Soc. Biol., Paris, 121, 1936, 408; Treponema ambigua Prdvot, Man. Class, et Determ. d. Bact^ries Ana^robies, Paris, 1940, 208.) From the oral cavity and the lungs. Pathogenic. Strict anae- robe . Spirochaeta amphibiae Yakimoff and Miller. (Bull. Soc. Path. Exot., 18, 1925, 306.) From the intestines of frogs, Rann temporaria . Spirochaeta argcnliucnsis Kuhn and Steiner. (Kuhn and Steiner, Med. Klin., 13, 1917, 1007; Spirochaeta poly- sclerotica Arzt and Kerl, according to Pettit, Contribution a I'Etude dcs Spiro- ch^tides, Vanves, II, 1928, 134; Trep- onema (?) argentinensis Xoguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 478.) From the livers of guinea pigs and rabbits inoculated with blood from patients liaving multiple sclerosis. Pathogenic for man, monkeys, dogs, rabbits and guinea pigs. Named for the Latin name of the town of Stras- bourg {Argentoralum) . Spirochaeta balanitidis Hoffmann and Prowazek. (Hoffmann and Prowazek, Cent, f . Bakt., I Abt., Orig., 41, 1906, 741 ; Spiroschaudinnia balanitidis Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404; Spironema balanitidis Park and Williams, Pathogenic Micro- organisms, 6th ed., 1917, 505; Treponema balanitidis Brumpt, Nouveau Traits de Medeeine, Paris, 4, 1922, 496.) From a case of balanitis. Spirochaeta bovis-caffris Nuttall. (Nuttall, Parasitology, 3, 1910, 108; Spironema bovis-caffris Ford, Textb. of Bact., 1927, 900.) From the blood of a buffalo . Spirochaeta bronchialis Castellani. (Castellani, Ceylon Medical Reports, 1907; Spiroschaudinnia bronchialis Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 402; Treponema bronchiole Brumpt, Nouveau Traite de Medeeine, Paris, 4, 1922, 496.) Found in cases of bronchitis in Ceylon. A mixture of sev- eral species of mouth spirochaetes is ap- parently described under this designation. Spirochaeta bucco-pharyngei Macfie. (Macfie, Ann. Trop. Med. and Parasitol., 10, 1916, 329; Treponema bucco-pharyngei Brumpt, Nouveau Traite de Medeeine, Paris, 4, 1922, 497.) From the throat of a native of the Gold Coast. May be identical with Spirochaeta dentium or S. buccalis. 1066 MANUAL OF DETERMINATIVE BACTERIOLOGY Spirochaeta bufonis Dobell. (Dobell, Quart. Jour. Microsc. Sci., 52, 1908, 121; Spiroschaudinnia hiijonis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 454 ; Spironema bufonis Ford, Textb. of Bact., 1927, 986; Treponema bufonis Ford, ibid.) From the intestines of a toad, Bufo vulgaris. Spirochaeta caesirae relortifurniis Hell- mann. (Arch. f. Protistenk., 29, 1913, 22.) From the urinary sac of a tunicate, Caesira retortiformis. Spirochaeta caesirae septentrionalis Hellmann. (Arch. f. Protistenk., 29, 1913, 22.) From the urinary sac of a tunicate, Caesira septentrionalis. Spirochaeta canina Bosselut. (Bull. Soc. Path. Exot., 18, 1925, 702.) From the blood of a dog. Spirochaeta canis Macfie. (Ann. Trop. Med. and Parasitol., 10, 1916, 305.) From dog feces. Spirochaeta cobayae Knowles ami Basu (Knowles and Basu, Indian Jour. Med. Res., 22, 1935, 449; Treponema cobayae Topley and Wilson, Princip. Bact. and Immun., 2nd ed., 1936, 725; Borrelia cobayae Steinhaus, Insect Microbiology, 1946, 454.) From the blood of guinea pigs. Blood parasite belonging to the relapsing fever group. Pathogenic for guinea pigs, rabbits and white rats. Spirochaeta comandoni Seguin and Vinzent. (Seguin and Vinzent, Compt. rend. Soc. Biol., Paris, 121, 1936, 408; Treponema comandoni Prevot, Man. Class, et Determ. d. Bact^ries Anaero- bies, Paris, 1940, 208.) From the oral cavity. Rather common. Non-patho- genic. Strict anaerobe. Spirochaeta crocidurae Leger. (Leger, Bull. Soc. Path. Exot., 10, 1917, 280; Treponema crocidurae McFarland, Patho- genic Bacteria and Protozoa, 2nd ed., 1933, 136.) From a shrew-mouse, Croci- dura stampjiii, in Senegal. Transmitted by Ornithodoros erraticus. Spirochaeta ctenocephali Patton. (Patton, Ann. Trop. Med. and Parasitol., 6, 1912, 357 ; Treponema ctenocephali Ford, Textb. of Bact., 1927, 989.) Para- sitic in the digestive tract of the larvae of the Indian cat-flea, Ctenocephalus felis . Spirochaeta cubensis Hoffman. (Sani- dad y Beneficienca Bolctin oficial, Ha- vana, 28, 1923, 76.) From the feces of Ilyla septentrionalis . Spirochaeta culicis JsiHo. (Jaff^, Arch, f. Protistenk., 9, 1907, 100; Spironema culicis Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 87; Entomospira culicis Endei'lein, Sitzungsber. Ges. Naturf. Freunde, Berlin, 1917, 313; Spiro- schaudinnia culicis Castellani and Chal- mers, Man. Trop. Med., 3rd ed., 1919, 454; Spirillum culicis Pringault, Compt. rend. Soc. Biol., Paris, 84, 1921, 209; Treponema culicis Ford, Textb. of Bact., 1927, 989.) Found in the intestines and Malpighian tubules of mosquito larvae, Culex sp. Spirochaeta didelphis Vianna, de Figu- eiredo and Cruz. (Brasil-Medico, 26, 1912, 912.) From the blood of an opos- sum, Didelphis aurita. Spirochaeta cqui (Novy and Knapp) Castellani and Chalmers. (Spirillum equi Novy and Knapp, Jour. Inf. Dis", 3, 9106, 294; Castellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 309; Spiroschaudinnia equi Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404; Spironema equi Noguchi, Jour. Exp. Med., 27, 1918, 584; Treponema equi Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 461.) From the blood of a horse. May be identical with Borrelia theileri. , Spirochaeta equina. (Dodd ?, Jour. Comp. Pathol, and Therap., 19, 1906, 318; quoted from Pettit, Contribution a I'Etude des Spirochetid^s, Vanves, II, 1928, 111.) Spirochaeta eurygyrata Werner. (Wer- ner, Cent. f. Bakt., I Abt., Grig., 52, 1909, 241; Spironema curgyratum (sic) Noguchi, Jour. Exp. Med., 27, 1918, 584; Spiroschaudinnia eurygyrata Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 451 ; Spirillum eurygyrata Castel- lani and Chalmers, ibid.; Borrelia eury- FAMILY TREPONEMATACEAE 1067 gyrata Brumpt, Nouveau Traits de Mede- cine, Paris, 4, 1922, 495; Treponema eurygyratum Brumpt, ibid., 50.) From the intestinal contents of man. Spirochaeta exanthematotyphi Futaki. (Futaki, Brit. Med. Jour., Oct. 13, 1917; Treponeyna exanthemalotyphi Sa- vini, Compt. rend. Soc. Biol., Paris, 88, 1923, 958.) Found in the kidneys and urine of cases of exanthematous typhus. Non -pathogenic . Spirochaeta febris Chester. (Afaiias- siew, Cent. f. Bakt., I Abt., 25, 1899, 405; Chester, Man. Determ. Bact., 1901, 347.) From a case of recurrent fever. Spirochaeta gadi Neumann. (Neu- mann, Ztschr. f. Hyg., 64, 1909, 79; Microspironona gadi Duboscq and Le- bailly, Compt. rend. Acad. Sci. Paris, 154, 1912, 662; Treponema gadi Duboscq and Lebailly, Arch. zool. exper. et gener., 10, 1912, 331; Spironema gadi Ford, Textb. of Bact., 1927, 964.) From the blood of a sea fish, Gadus minutus. Spirochaeta gallica Couvy and Dujar- ric de la Riviere. (Couvy and Dujarric de la Riviere, Compt. rend. Soc. Biol., Paris, 81, 1918, 22; Treponema gallicum Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 500.) From the blood of trench fever patients. Spirochaeta gangraenae carcinomatosae Hoffmann. (Berl. klin. Wochnschr., 42, 1905, 880.) From malignant tumors. Spirochaeta gangraenosa nosocomialis Rdna. (Rona, Verhandl . d. deutsch . der- mat. Gesellsch., 9, 1907, 471; Treponema gangraenosa nosocomialis Noguchi, Jour. Exp. Med., 16, 1912, 261.) From ulcers of the genital region. Spirochaeta haemophilus Trosier and SifTerlen. (Ann. Inst. Past., 58, 1937, 233.) From a child with intestinal trou- ble and continuous fever. Spirochaeta hispanica de Buen. (De Buen, Ann. Parasitol., 4, 1926, 185; Trep- onema hispanicum Nogucjii, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 481; Spirochaeta maro- canum Nicolle and Anderson, Compt. rend. Acad. Sci. Paris, 187, 1928, 747; Spirochaeta hispanicum \ar. niarocaniun Nicolle, Anderson and Colas-Belcour, Arch. Inst. Past. Tunis, 18, 1929, 343; Treponema hispanicum var. marocanum Gay et al.. Agents of Disease and Host Resistance, 1935, 1074; Borrelia his- panicum Steinhaus, Insect Microbiology, 1946, 453.) The cause of Spanish and INIoroccan relapsing fever. Transmitted by Ornithodoros marocanus. Not ag- glutinated by serum of Borrelia recur- rentis. Pathogenic for man and laboratory animals. Spirochaeta intestinalis Macfie and Carter. (Macfie and Carter, Ann. Trop. Med. and Parasitol., 11, 1917, 79; Trep- onema intestinale Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 505.) From human feces. Spirochaeta jonesii Dutton, Todd and Tobey. (Dutton, Todd and Tobey, Jour. Med. Res., 15 (N.S. 10), 1906, 491; Spironema jonesii Ford, Textb. of Bact., 1927, 964.) From the blood of an African mudfish, Clarias angolensis. Spirochaeta lagopodis Fantham. (Fan- tham, Proc. Zool. Society, London, 1910, 692; Spironema lagopodis Noguchi, ac- cording to Pettit, loc. cit.) From the blood of the grouse, Lagopus scoticus. Spirochaeta leucotermitis HoUande. (Arch. zool. exper. et g6n., 61, 1922, 23.) From an insect, Leucotermes lucif^tgus. Spirochaeta lovati Fantham. (Proc. Zool. Society, London, 1910.) From the intestinal contents of the grouse, Lagopus scoticus. Spirochaeta lowenthali Besson. (Bes- son, p. 736, according to Ford, Textb. of Bact., 1927, 1001.) From malignant tumors. Spirochaeta lutrae Prowazek. (Pro- wazek, Arb. kaiserl. Gesundheitsamte, 26, 1907, 31 ; Spiroschaudinnia lutrae Cas- tellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404 ; Spironema lutrae Ford, Textb. of Bact., 1927, 961.) From the blood of an otter, Lutra sp. Spirochaeta lymphaticus Proescher and White. (Proescher and White, Jour. Amer. Med. Assoc, 49, 1907, 1988; 1068 MANUAL OF DETERMINATIVE BACTERIOLOGY Treponema lijmphaiicum Brumpt, Nou- veau Traits de M^decine, Paris, 4, 1922, 506.) From cases of lymphatic leukemia. Pathogenic for guinea pigs, rats and monkeys. Spirochaeta macaci Castellani and Chalmers. (Castellani and Chalmers, 1908, see Castellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 310; Spiro- schaudinnia macaci Castellani and Chal- mers, Man. Trop. Med., 2nd ed., 1913, 403; Spironema macaci Ford, Textb. of Bact., 1927, 960.) From monkeys in Cey- lon. Closely related to or possibly itlen- tical with Borrelia carteri. Spirochaeta marmoiae Carpano. (Ann . d'Igien. sper., 23, 1913, 215.) From a rodent, Arctomys marmota. Spirochaeta titelanogenes canis Lukes. (Deutsch. Tierarztl. Wochnschr., 1923, 137; see Ann. Inst. Past., 38, 1924, 523.) From the intestines of a dog. The cause of a dog plague . Spirochaeta melophagi Porter. (Quart . Jour. Microsc. Sci., London, 55, 1910, 189.) From the intestine, ovaries, and pupa of Melophagus ovinus. Spirochaeta inetritis de Andrado. (Compt. rend. Soc. Biol., Paris, 86, 1922, 1048.) From a uterine excretion. Spirochaeta microgyrata gaylordi Cal- kins. (Calkins, Jour. Inf. Dis., 4, 1907, 173; Spirochaeta microgyrata var. gaylordi Calkins, ibid., 171; Spironema microgy- rata var. gaylordi Ford, Textb. of Bact., 1927, 1001 . ) From breast tumors of mice . Spirochaeta naganophila Savini. (Compt. rend. Soc. Biol., Paris, 88, 1923, 956.) From the blood of mice. No description given. Spirochaeta noelleri Zuelzer. (1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1684.) From the intestine of the larvae of Simulium noelleri. Spirochaeta noguchii Strong. (Strv)ng, United Fruit Co. Med. Dept. 14th Ann. Kept., 1925; Treponema noguchii Nogu- chi, in Jordan and Falk, Newer Knowl- edge Bact. and Immun., 1928, 483.) From a type of skin ulcer in South America. Spirochaeta normandi Nicolle, Ander- son and Colas-Belcour. (Compt. rend. Acad. Sci. Paris, 185, 1927, 334.) From Ornithodoros normandi, the natural agent of transmission. Pathogenic for white mice . Spirochaeta normandi var. carthaginen- •sis. (Quoted from Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 169.) From ticks in Tunis. Spirochaeta nosocomialis Hoffmann and Gonder. (Hoffmann anrl Gonder, 1914, according to Brumpt, Nouveau Trait(5 de M^decine, Paris, 4, 1922, 508; Treponeiria nosocomiale Brumpt, ibid.) Probably a synonym of Borrelia vincentii. Spirochaeta obtusa Castellani. (Cas- tellani, Brit. Med. Jour., 3, 1905, 1330; Spirochaeta tenuis obtusa Castellani, 1908, according to Ford, Textb. of Bact., 1927, 973; Spiroschaudinnia obtusa Cas- tellani, Arch. f. Schiffs- u. Tropenhyg., 12, 1908,311 (see Castellani and Chalmers, Man.. Trop. Med., 3rd ed., 1919 449); Treponema obtusum Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 508; Treponema tenve cbtusum Brumpt, ibid.) From ulcerated lesions of yaws. Spirochaeta ovis (Novy and Knapp) Bergey ct al. {Spirillum ovis Novy and Knapp, Jour. Inf. Dis., 3, 1906, 294; Spirochaeta ovina Blanchard, Semaine M^dicale, 28, 1906, 1; Spiroschaudinnia ovina Castellani and Chalmers, Man. Trop. Med., 2nd ed., 1913, 404; Spiro- nema ovis Ford, Textb. of Bact., 1927, 960; Treponema ovis Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 461; Treponema ovinum Noguchi, ibid., 480; Bergey et al.. Man- ual, 5th ed., 1939, 960.) From the blood of sheep. May be identical with Borrelia theileri . Spirochaeta pelamidis Neumann. (Neumann, Ztschr. f. Hyg., 64, 1909, 80; Spironema pelamidis Ford, Textb. of Bact., 1927, 964.) From the blood of a fish, Pelamys sarda. Resembles Spiro- chaeta gadi. Spirochaeta perforans Cavalie and Man- doul. (Compt. rend. Soc. Biol., Paris, FAMILY TREPONEMATACEAE 1069 85, 1921, 1068.) From cases of pyorrhea alveolaris. Associated with fusiform ba- cilli. Probably synonj-mous with Bor- relia vinceyitii. Spirochaeta periplanetae Laveran and Franchini. (Bull. Soc. Path. E.xot., 13, 1920, 332.) From the digestive tract of the cockroach, Pcriplaneta orientalis. Spirochaeta pcrsica Dschunkowsky. (Dschunkowsky, Deutsch. med. Woch- nschr., 39, 1913, 419; Treponema persi- cum Brumpt, Xouveau Traite de Mede- cine, Paris, 4, 1922, 509; Borrelia persica Steinhaus, Insect Microbiology, 1946, 452.) From a case of relapsing fever (Mianeh fever) in Persia. Transmitted by Ornithodoros tholozani and 0. latio- rensis. Serum not agglutinated by Borrelia recur reniis. Disease in man fairly severe and in gerbilles and mon- keys very mild. Spirochaeta pollachii Henry. (Spiro- chaeta gadi pollachii Henry, Jour. Path, and Bact., 14, 1910, 463; Henry, ibid., 17, 1912, 160; Treponema fallaxDuhoscq and Lebailly, Arch. Zool. Exper. et Gen., 10, 1912, 331 ; Spirochaeta fallax Zuelzer, 1925, in Prowazek, Handb. d. path. Pro- toz., S, 1931, 1671; Spironema pollachii Ford, Textb. of Bact., 1927, 965.) From the blood of the pollack, Gadtis polla- chius . Spirochaeta pseudobuccalis Zuelzer. (Cent. f. Bakt., I Abt., Orig., 85, 1921, *154.) Spirochaeta pseudorecurrentis Zuelzer. (Cent. f. Bakt., I Abt., Orig., 85, 1921, *154.) Spirochaeta pyorrhoeica Kolle. (Med. Klin., 13, 1917, 59.) Commonly found in pus from pyorrhea alveolaris. Spirochaeta raillieti Mathis and Leger. (Compt. rend. Soc. Biol., Paris, 70, 1911, 212.) From the blood of a rabbit. Not pathogenic . Spirochaeta ranarum Yakimoff and Miller. (Bull. Soc. Path. Exot., 18, 1925, 306.) From the intestines of frogs, Rana temporaria. Spirochaeta recta Gerber. (Gerber, Cent, f . Bakt., I Abt., Orig., 56, 1910, 513 ; Treponema rectum Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922,511.) May be a synonym of Borrelia vincentii. Spirochaeta regaudi Ball and Roquet. (Ball and Roquet, 1911, according to Pettit, Contribution a I'Etude des Spiro- ch^tides, Vanves, II, 1928; Spirella regaudi Ball and Roquet, 1911, according to Brumpt, Xouveau Traits de IMedecine, Paris, 4, 1922, 517; also see Edkins, Parasitology, 15, 1923, 296.) From the stomachs of cats and dogs. Possibly be- longs among the spirilla (Noguchi) ; is in the same group as Cristispirella and Heliconema. Spirochaeta sinensis Pons. (Compt. rend. Soc. Biol.; Paris, 89, 1923, 1028.) From the blood of a fever patient in China. Pathogenic for rabbits and mon- keys. Spirochaeta sogdianum Nicolle and Anderson. (Nicolle and Anderson, Compt. rend. Acad. Sci. Paris, 187, 1928, 746; Borrelia sogdianum Steinhaus, Insect Microbiology, 1946, 453.) Not pathogenic for guinea pigs or fowls. Probably synonymous with Borrelia recurrentis. Spirochaeta sporogenes psoriasis Rasck. (Individual publications, Christiania, 1920-1921, 4.) Spirochaeta sporogona rheumatismi Rasck. (Individual publications, Chris- tiania, 1920-1921, 4.) From the blood in cases of acute arthritis. Spirochaeta staphylina Ghidini and Archetti. (Riv. Biol. Coloniale, 2, 1939, 131.) From the intestine of a termite, Reticulitermes lucifugus, Italy. Spirochaeta stenogyrata Werner., (Werner, Cent. f. Bakt., I Abt., Orig., 52, 1909, 241; Treponema stenogyratum Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922, 514.) From human feces. Spirochaeta suilla Dodd. (Jour. Compt. Path, and Therap., 19, 1906, 216.) From cutaneous lesions of pigs. Patho- genic for pigs. Spirochaeta temporariae Yakimoff and Miller. (Bull. Soc. Path. Exot., 18, 1070 MANUAL OF DETERMINATIVE BACTERIOLOGY 1925, 306.) From the intestines of frogs, Rana temporaria. Spirochaeta tenuis Gerber. (Gerber, Cent. f. Bakt., I Abt., Orig., 56, 1910, 508; Treponema tenue Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922, 514.) May be identical with Spirochaeta den- tium or with Borrelia vincentii. Spirochaeta termitis (Leidy) Dobell. {Vibrio termitis Leidy, Jour. Acad. Nat. Sci., Phila., 2nd Ser., 8, 1881, 441 ; Spiro- chaeta minei Prowazek, Arch. f. Schiffs- u. Tropenhyg., U, 1910, 297; Dobell, Spolia ceylanica, 3, 1910, 78; Treponema minei Dobell, ibid.; ? Spirochaete grassii Doflein, Prob. der Protistenk., 2, 1911, 17; Spirochaeta grassii Doflein, Die Natur der Spirocliaten, Jena, 1911; Treponema tenniiis Dobell, Arch. f. Protistenk., 26, 1912, 117; Entomospira grassii Enderlein, Sitzungsber. Ges. Naturf. Freunde, Berlin, 1917, 313; Cristispira termitis Hollande, Arch. Zool. Exper. et G^n., 61, 1923, N. and R., 25; Treponema grassi Ford, Textb. of Bact., 1927, 988.) From the intestines of Termes lucifugus and Calotennes spp. Spirochaeta tropidonoti Dobell. (Do- bell, Spolia ceylanica, 7, 1911, 65; Spiro- nema tropidonoti Ford, Textb. of Bact., 1927, 962.) Isolated once from the blood of a snake, Tropidonotus stolatus, in Ceylon. Spirochaeta urethrae IMacfie. (Macfie, Ann. Trop. Med. and Parasitol., 10, 1916, 305; Spiroschaudinnia urethrae Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 451; Treponema urethrae Brumpt, Nouveau Traite de Medocine, Paris, 4, 1922, 514.) From the urine of Gold Coast natives. Causes acute arthritis. Spirochaeta u^bekistanica Pickoul. (Russ. Jour. Trop. Med., 6, 1928, 612.) From cases of relapsing fever in Bokhara. Spirochaeta vespertilionis (Novy and Knapp) Castellani and Chalmers. (Spi- rillum vespertilionis Novy and Knapp, Jour. Inf. Dis., 3, 1906, 294; Castellani and Chalmers, Man. Trop. Med., 1st ed., 1910, 309; Spiroschaudinnia vespertilionis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 454; Spironema ves- pertilionis Ford, Textb. of Bact., 1927, 961.) From the blood of a bat, Vesper- tilio kuhlii. Spirochaeta vincenti var. bronchialis Delamare. (Compt. rend. Soc. Biol., Paris, 50, 1924,611.) Spirochaeta zlatogorori Yakimoff. (Bull. Soc. Path. Exot., 14, 1921, 532.) From feces. Spirochaete exanthematica Lewascheff. (Cent. f. Bakt., I Abt., 18, 1895, 133.) From the blood in cases of typhus fever. Spirochaete forans Reiter. (Reiter, Deutsch. med. Wochnschr., No. 50, 1916, 10; see Cent. f. Bakt., I Abt., Orig., 79, 1917, 176; Treponema forans Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 500; Spirochaeta forans Pettit, Con- tribution a I'Etude des Spirochetides, Vanves, II, 1928, 164.) From the l)lood in a case of articular rheumatism. Not pathogenic for guinea pigs or mice. Spirochaete gracilis Veszpremi. (Veszpremi, Cent. f. Bakt., I Abt., Orig., 44, 1907, 332; not Spirochaeta gracilis Levaditi and Stanesco, Compt. rend. Soc. Biol., Paris, 67, 1909, 188 (Treponema levaditii Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 501 ; Treponema gracile Brumpt, idem); Treponema gra- cile Ford, Textb. of Bact., 1927, 978.) From a gangrenous phlegmon of the mouth. Found in association with fusi- form bacilli and therefore may be identi- cal with Borrelia vincentii or Spirochaeta dentium or Treponema macrodentium . Spirochaete repacis. (Quoted from Lehmann and Neumann, Bakt. Diag., 6 Aufl., ;2, 1920, 809.) From the oral cavity. Spironema caviae Ford. (Spirochaete (?), Macfie, Ann. Trop. Med. and Para- sitol., 8, 1914, 447; Ford, Textb. of Bact., 1927, 961.) From the blood of a guinea pig at Lagos. Spironema vesper uginis (Gonder) Ford. (Spirochaeta vesperuginis Gonder, Arb. kais. Gesundheitsanite, 27, 1908, 406; Ford, Textb. of Bact., 1927, 961.) From the blood of a bat, Vesperugo kuhlii . Spiroschaudinnia caviae Sangiorgi. FAMILY TREPONEMATACEAE 1071 (Sangiorgi, Pathologica Rivista, 5, 1913, 428 ; Spirochaeta caviae Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 174.) From the blood of a guinea pig. Spiroschaudinnia luitis Castellani and Chalmers. (Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 451; Treponema mite Brumpt, Nouveau Traite de Medecine, Paris, 4, 1922, 506.) From urine in mild cases of camp jaundice. Probably not pathogenic. Treponema lineola (Donne) Brumpt. {Vibrio lineola Donne, Recherches Mi- crosc. Nature d. Mucus des Organs Geni- tourinaires, Paris, 1837; Brumpt, Nou- veau Traite de Medecine, Paris, 4, 1922, 505.) From secretions of the genitalia. Genua II. Treponema Schaudi7in. (Spironema Vuillemin, Compt. rend. Acad. Sci. Paris, I40, 1905, 1567 ; not Spironema Bergey et al., Manual, 1st ed., 1923, 424; Schaudinn, Deutsche med. Wochnschr., 31, 1905, 1728; Microspironema Stiles and Pfender, Amer. Med., 10, 1905, 936.) Length 3 to 18 microns. Longer forms due to incomplete division. Protoplasm in acute, regular or irregular spirals. Terminal filament may be present. Some species stain only with Giemsa's stain. Weakly refractive by dark field illumination in living preparations. Cultivated under strictly anaerobic conditions. Pathogenic and parasitic for man and animals. Generally produce local lesions in tissues. The type species is Treponema pallidum (Schaudinn and Hoffmann) Schaudinn. 1. Treponema pallidum (Schaudinn and Hoffmann) Schaudinn. {Spirochaete pallida Schaudinn and Hoffmann, Arb. kaiserl. Gesundheitsamte, 22, 1905, 528; Schaudinn, Deutsche med. Wochnschr., 31, 1905, 1728; Spironema pallidum Vuil- lemin, Compt. rend. Acad. Sci. Paris, I40, 1905, 1567; Microspironema pallidum Stiles and Pfender, Amer. Med., 10, 1905, 936; Trypanosoma luis Krzystalowicz and Siedlecki, 1905, see abst. in Bull. Inst. Past., 4, 1906, 204; Spirochaeta pallida Hoffmann and Prowazek, Cent, f. Bakt., I Abt., Orig., 4I, 1906, 741.) From Latin, pale. Morphology : Very fine protoplasmic spirals 0.25 to 0.3 by 6 to 14 microns. Spiral amplitude: 1.0 micron, regular, fixed. Spiral depth : 0.5 to 1.0 micron. Terminal spiral filament present. Weakly refractive in living state by dark field illumination. May appear as a series of bright dots or string of radiant beads with poor dark field illumination. Staining: Stain with difficulty except with Giemsa's stain by which they appear pink or rose. Appear black with silver impregnation methods. Motility: Sluggish, drifting motion, stifflj' flexible, rarely rotating. Trypsin digestion : Resistant for many days . Bile salts (10 per cent) : Disintegration complete. Saponin (10 per cent): Broken up in time. Cultivation: With difficulty under strict anaerobiosis in ascitic fluid with addition of fresh rabbit kidney. Habitat : The cause of syphilis in man. Can be transmitted experimentally to anthropoid apes and rabbits. 2. Treponema pertenue Castellani. (Castellani, Jour. Trop. Med., 8, 1905, 253; Spirochaeta pertenuis Castellani, Jour. Ceylon Branch Brit. Med. Assoc, June, 1905; Spirochaeta pallidula Castel- lani, Brit. Jour. Med., 2, Nov., 1905, 1330; Spirochaete pertemds Lehmann and Neu- mann, Bakt. Diag., 5 Aufl., 2, 1912, 677; Spironema pertenue Gross, Archiv f. Protistenk.,^4, 1912, 115; Treponemapal- lidulum Brumpt, Nouveau Traits de Medecine, Paris, 4, 1922, 508.) From Latin, very fine. 1072 MANUAL OF DETERMINATIVE BACTERIOLOGY Morphologically indistinguishable from Treponema pallidum. Cultivable under anaerobic conditions in the same medium used for Treponema pallidum. Habitat : The cause of yaws — tropica frambesia. Patients with the disease give a positive Wassermann test. Prbo- ably transmitted by contact. 3. Treponema microdentium Noguchi. (Jour. Exp. Med., 15, 1912, 81.) From Greek viikros, small and Latin, teeth. The organism is less than 0.25 micron in thickness in the middle and tapers toward each extremity, which is pointed. The length varies with age but may reach 8 microns and show an average of 14 curves. Sometimes a long, thin flagella-like pro- jection is observed at each extremity. Growth occurs under anaerobic condi- tions in serum water medium containing fresh tissue. The serum is slightly coag- ulated and gives off a strong, fetid odor. Habitat : Normal oral cavity. 4. Treponema mucosimi Noguchi. (Jour. Exp. Med., 16, 1912, 194; Spiro- chaeta mucosa Pettit, Contribution a I'Etude des Spirochetides, Vanves, II, 1928, 190.) From Latin, mucous. Spirals: 0.25 to 0.3 by 8 to 12 microns. The number of curves varies from 6 to 8. Both extremities are sharply pointed and often possess a minute curved projection, 8 to 10 microns long. Cultivable under anaerobic conditions, forming mucin. The cultures give off a strong, putrid odor. Takes the red in Giemsa's stain. Strict anaerobe. Source : From pus in a case of pyorrhoea. Habitat : Found in pyorrhea alveolaris. It possesses pyogenic properties. 5. Treponema calligyrum Noguchi. (Noguchi, Jour. Exp. Med., 17, 1913, 96; Spirochaeta calligyra Zuelzer, 1925, in Prowazek, Hand. d. path. Protoz., 3, 1931, 1673.) From M. L., with beautiful circles. Morphology: 0.35 to 0.4 by 6 to 14 mi- crons, average 9 to 12 microns. Spirals are regular and deep but more rounded than those of Treponema pallidum. The organism is of uniform width until near the extremities which end in sharp points with delicate projections. Motility : Active, chiefly rotating. Stains reddish-violet with Giemsa's stain. Cultivation: Grows under anaerobic conditions. Not pathogenic for monkeys or rabbits. Source: From smegma. Habitat : I^esions and membranes of the pudenda. 6. Treponema genitalis Noguchi. {Treponema minutum Noguchi, Jour. Exp. Med., 27, 1918, 671 ; not Treponema minutum Dobell, Arch, f . Protistenk., 26, 1912, 151 ; not Treponema minutum Castellani, 1916 ; Noguchi, Laboratory Di- agnosis of Syphilis, New York, 1923, 260; Spirochaeta minutum Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1673; Spirochaeta genitalis Seguin and Vinzent, Ann. Inst. Past., 61, 1938, 255.) From Latin, genital. Morphology: 0.25 to 0.3 by 3 to 14 mi- crons. Spirals round, regular and shal- low. Smaller than Treponema pallidum and spirals are closer together. Motility: Active. Culture : Grows anaerobically and re- quires fresh tissue. Non-pathogenic . Habitat: Found on mule and female genitalia. 7. Treponema carateum Brumpt. (Treponema de un caso de pinta, Saenz, Grau Triana and Alfonso, Arch, de Med. Int., Havana, 4, 1938, 3; Brumpt, Compt. Rend. Soc. Biol., Paris, 130, 1939, 942; Treponema herrejoni Leon y Blanco, Rev. de Med. Trop. y Parasitol., Habana, 6, 1940, 5; Treponema pictor Pardo-Cas- tello, Rev. de Med. Trop. y Parasitol FAMILY TREPONEMATACEAE 1073 Habana, 6, 1940, 117; Treponema pinlae Curbelo, Elementos de Bacteriologia Medica, 1941, 34.) From carate, spotted sickness. Description taken from Leon y Blanco {loc. cit.). Cylindrical: 0.25 to 0.30 by 7.8 to 36.8 microns, average length 17.8 microns. With sharp-pointed ends. Spiral amplitude : 1 micron, regular. Spiral depth: 0.8 to 1.0 micron. Number of waves, 6 to 27, according to length. Ten to twelve (Brumpt, loc. cit.). Actively motile. At times undulating or creeping movements are shown. Staining reactions : Readily takes silver impregnations, Giemsa's stain, carbol- fuchsin and gentian violet. Saponin (10 per cent) : Disintegrates in six hours at room temperature. Same result with sodium taurocholate (10 per cent) and with bile. Distilled water : Produces swelling. Loses motility on heating for 15 minutes at 50°C or for 3 hours at 4rC. Wassermann, Kahn and Meinicke reac- tions positive. Has not yet been cultivated artificially. Experimental transmission unsuccessful so far. Source : From the border of cutaneous lesions of persons having pinta (spotted sickness). Habitat : The cause of pinta (or carate) . Common in Mexico and Colombia. Also found in other northern countries of South America, in Central America and the West Indies. Rare in Cuba. Pos- sibly found in other tropical regions of the world. 8. Treponema cuniculi Noguchi. {Spirochaeta paraluis cuniculi Jakobs- thal, Dermatol. Wchnschr., 71, 1920, 569; Noguchi, Jour. Amer. Med. Assoc, 77, 1921, 2052; also see Noguchi, Jour. Exp. Med., 35, 1922, 395; Treponema pallidum var. cuniculi Klarenbeek, Cent. f. Bakt., I Abt., Orig., 87, 1921, 203; Spirochaeta cuniculi Seraditi, Marie and Isaien, Compt. rend. Soc. Biol., Paris, 85, 1921, 51 ; Spirochaeta pallida var. cuniculi Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1765; Spirochaeta paraluis Pettit, Contribution a I'Etude des Spirochetides, Vanves, II, 1928, 91; Spirochaeta paraluis-cuniculi Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 187.) From Latin, rabbit. Description from Noguchi (loc. cit.). Closely resembles Treponema pallidum, but longer. Width 0.25 micron; length 10 to 16 mi- crons; long specimens up to 30 microns frequent. Spirals 8 to 12 in number, regular, deep. Spiral amplitude 1 to 1.2 microns. Spiral depth 0.6 to 1.0 micron. Delicate terminal filament at one, sometimes both, ends. Often forms entangled masses of long threads; occurs sometimes in a stellate arrangement . Staining properties same as for Trepo- nema pallidum. Both readily stained by ordinary basic analine dj'es when fixed in a buffered formaldehyde solution. Wassermann reaction negative. Pathogenesis : Disease transmissible to healthy rabbits, producing papular le- sions in the genitoperineal region. Not pathogenic for monkeys, mice or guinea pigs. Source : From lesions in the genitoperi- neal region of five rabbits. Habitat : The cause of rabbit spiro- chetosis. Appendix: Many of the species in this appendix are so inadequately described that it is not certain that they belong in this group. Microspironema legeri Duboscq and Lebailly. (Duboscq and Lebailly, Compt. rend. Acad. Sci. Paris, 154, 1912, 662; Treponema legeri Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz. , 3, 1931, 1683.) From a fish. Box hoops. Spirochaeta microgyrata Loewenthal. (Loewenthal, Berl. klin. Wochnschr., 4^, 1906, 283; Spironema microgyrata Nogu- 1074 MANUAL OF DETERMINATIVE BACTERIOLOGY chi, Jour. Exp. Med., ^7, 1918, 584 ; Spiro- schaudinnia microgyrata Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 454 ; Treponema microgyratum Brumpt, Nouveau Traits de M^decine, 4, 1922, 506.) From cancerous ulcers of man, dogs, and mice. Regarded by Hoff- mann and Prowazek (Cent. f. Bakt., I Abt., Orig., 4i, 1906, 819) as identical with Spirochaeia dentium. Spirochaeta parotitidis Lehmann. (In Lehmann and Neumann, Bakt. Diag., 7 Aufl., 2, 1927, 580.) Pathogenic, produc- ing a disease similar to mumps in experi- mental animals (cats) and causing paro- titis and orchitis in apes. Spirochaeta penortha Beve ridge. (Aus- tral. Jour. Expt. Biol, and Med. Sci., 14, 1936, 307.) This organism is also called Treponema podovis according to Shahan, in Keeping Livestock Healthy, U. S. D. A. Yearbook of Agriculture, Part 6, 1942, 830.) Present as an accessory fac- tor in foot-rot of sheep. Also see Acti- nomyces nodosus. Spirochaeta phlebotomi Pringault. (Compt. rend. See. Biol., Paris, 84, 1921, 209.) From the sand fly, Phlehotomus perniciosus. Spirochaeta pseudopallida Mulzer. (Mulzer, Berl. klin. Wchnschr., 42, 1905, 1144; Spiroschaudinnia pseudopallida Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919,449; Treponema pseu- dopallidum Brumpt, Nouveau Traits de MMecine, Paris, 4, 1922, 511.) Found in ulcerating carcinomata. Spirochaeta skoliodonta Hoft'mann. (Hoffmann, Cent. f. Bakt., I Abt., Orig., 86, 1920, 137 ; Spirochaeta acuta Kritchev- sky and S^guin, 1920, according to Seguin and Vinzent, Ann. Inst. Past., 67, 1941, 62; Treponema skoliodontum Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 481.) From the peri- toneal exudate of a guinea pig. One of the smallest known spirochetes. From the oral cavity. Spirochaeta subtilis Castellani. (Cas- tellani, 1907; Spiroschaudinnia subtilis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 450; Treponema sub- tile Brumpt, Nouveau Traits de M^de- cine, Paris, 4, 1922, 514; Spironema sub- tilis Pettit, Contribution k I'Etude des Spiroch^tid^s, Vanves, II, 1928.) From the oral mucosa and from intestinal con- tents. May be a synonym of Spirochaeta dentium. Spirochaeta urethralis Castellani. (Castellani, 1915; Treponema urethrale Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 1944.) From a puru- lent urethral discharge. Spirochaeta vaccinae Bonhoff. (Bon- hoff, Berl. klin. Wochnschr., 42, 1905, 1142; Treponema vaccinae Brumpt, Nou- veau Traits de M^decine, Paris, 4, 1922, 515.) From vaccinia. Spirochaeta vaginalis Macfie. (Mac- fie, Ann. Trop. Med. and Parasitol., 10, 1916, 315; Treponema vaginalis Brumpt, Nouveau Traits de Mddecine, Paris, 4, 1922, 514.) From a case of vaginitis in a Gold Coast native. Spirochaete hartmanni Gonder. (Gon- der. Cent, f . Bakt., I Abt., Orig., 47, 1908, 491 ; Spironema hartmanni Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 88; Spiro- chaeta hartmanni Noguchi, Jour. Exp. Med., 34, 1921, 297; Treponema hart- manni Ford, Textb. of Bact., 1927, 990.) From the digestive tract of molluscs. Pinna spp. Spirochaete polyspira Wolff. (Wolff, Cent. f. Bakt., II Abt., 18, 1907, 448; Treponema polyspirum Wolff, ibid.; Spi- rochaeta polyspira Pettit, Contribution k I'Etude des Spirochetid^s, Vanves, II, 1928, 14.) From rotten potatoes. Treponema carpanoi Yakimoff and Rastjapin. (Arch. f. Protistenk., 71, 1930, 543.) From stomatitis of horses. Treponema cotti Duboscq and Lebailly. (Duboscq and Lebailly, Arch. Zool. Ex- p^r. et G^n., 10, 1912, 331; Microspiro- nema cotti Duboscq and Lebailly, Compt. rend. Acad. Sci., Paris, 154, 1912, 662.) From the marine bullhead, Cottus btibalis. Treponema dentium (Miller) Dobell. (Spirochaete im Zahnschleim, Cohn, Beitr. z. Biol. d. Pflanzen, /, Heft 2, 1872, FAMILY TREPONEMATACEAE 1075 180; Spirochaete denticola Fliigge, Die Mikroorganismen, 2 Aufl., 1886, 390; Spi- rochaete dentium Miller, Microorganisms of the Human Mouth, Philadelphia, 1890, 80; Spirillum dentium Sternberg, Manual of Bact., 1893, 694; Spirochaeta dentium Migula, in Engler and Prantl, Die nattirl. Pflanzenfam., 1, la, 1895, 35; Spirochaeta denticola Arndt, according to Hoffmann and Prowazek, Cent. f. Bakt., I Abt., Orig., 41, 1906, 819; Dobell, Arch. f. Protistenk., 26, 1912, 117 ; Spironema den- tium Gross, Cent. f. Bakt., I Abt., Orig., 65, 1912, 88 ; Spirochaeta dentinum McFar- land. Pathogenic Bacteria and Protozoa, 7th ed., 1912, 546; Treponema microden- iium Noguchi, Jour. Exp. Med., 15, 1912, 81 ; Spirocfuieta orthodonta Hoffmann, Deutsch. med. Wochnschr., 46, 1920, 257; Spirochaeta microdentium Heim, Lehr. d. Bakt., 6 and 7 Aufl., 1922, 477 ; Treponema denticola Brumpt, Nouveau Traits de MMecine, Paris, 4, 1922, 497; Treponema orthodontum Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 481; Treponema dentium -sieno- gyratum Pettit, Contribution a 1 'Etude des Spiroch6tid6s, Vanves, II, 1928, 240.) The smallest of the mouth spirochaetes. Non-pathogenic. This term probably in- cludes several morphologically similar species which have not as yet been suffi- ciently characterized. Treponema drosophilae Chatton. (Compt. rend. Soc. Biol., 73, 1912, 212.) From Drosophila confusa. Six to thirty microns in length, tapers at both ends, four spirals, movement helicodal. Treponema gallicolum Lebailly. (Compt. rend. Soc. Biol., Paris, 75, 1913, 389.) From the caecum of the hen, Gal- lus sp. Treponema hilli Duboscq and Grasse. (Compt. rend. Soc. Biol., Paris, 94, 1926, 34; Arch. Zool. Exper. et Gen., 66, 1927, 484.) From the surface of the body of a flagellate, Devescovina hilli, and in the intestine of a termite, Glyplotermes iridipennis. A very small organism. Treponema intermedium Dobell. (Mittelformen, Liihe, Handb. d. Tropen- krankh., S, 1906; see Hoffmann and Pro- wazek, Cent. f. Bakt., I Abt., Orig., 41, 1906, 819 ; Dobell, Arch, f . Protistenk., ^&, 1912, 117; Treponema macrodentium No- guchi, Jour. Exp. Med., 15, 1912, 81; Spirochaeta media oris Hoffmann, Deutsch. med. Wochnschr., 46, 1920, 257 ; Treponema medium Brumpt, Nouveau Traits de M^decine, Paris, 4, 1922, 505; Spirochaeta intermedia Pettit, Contribu- tion a I'Etude des Spiroch^tid^s, Vanves, II, 1928, 146; Spirochaeta macrodentium Pettit, ibid., 182; Spirochaeta media and Spironema media Pettit, ibid., 240.) The middle-sized spirochete of the mouth. Treponema lari Lebailly. (Compt. rend. Soc. Biol., Paris, 75, 1913, 389.) Found in the caecum of birds, also in the guinea-pig. Named for one of the birds, Larus ridibundus. Treponema minutum Dobell. {Trepo- nema sp. Dobell, Quart. Jour. Microsc. Sci., 52, 1908, 121; Dobell, Arch. f. Pro- tistenk., 26, 1912, 151; not Treponema minutum Castellani, 1916; not Treponema minutum Noguchi, Jour. Exp. Med., 27, 1918, 671 ; Spirochaeta minutum Zuelzer, 1925, in Prowazek, Handb. d. path. Pro- toz., .3, 1931, 1682.) From the large intes- tines of toads, Bufo vidgaris. Treponema parvum Dobell. (Dobell, Arch. f. Protistenk., 26, 1912, 111 ; Spiro- chaeta parvum Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1685.) From the intestine of the cockroach, Sty- lopyga {Blatta, Periplaneta) orientalis. Very small organism. T'repojiema pavonis Duboscq and Le- bailly. (Arch. Zool. Expdr. et Gen., 10, 1912, 331.) From the intestine of the blenny, Blennius pavo. Treponema perexile Duboscq and Le- bailly. (Duboscq and Lebailly, Arch. Zool. Exper. et Gen., 10, 1912, 331 ; Spiro- chaeta perexilis Hindle, Med. Res. Coun- cil Syst. of Bact., 8, 1931, 180.) From the blood of a marine fish, Lepadogaster bimaculatus. Treponema podovis Ludovic and Blai- zot. (Compt. rend. Acad. Sci. Paris, 187, 1076 MANUAL OF DETERMINATIVE BACTERIOLOGY 1928, 911.) Pathogenic. Cause of a dis- ease in sheep. Treponema querquedulae Lebailly. (Compt. rend. Soc. Biol., Paris, 75, 1913, 389.) From caeca of birds. Named for the teal, Querquedula querquedula. Treponema rhinopharyngeum Brumpt. {Treponema minutum Castellani, 1916; Spiroschaudinnia minuta Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 1881; Brumpt, Nouveau Traite de Mddecine, Paris, 4, 1922, 514.) From man in cases of rhinopharyngitis. Prob- ably a synonym of Spirochaeta gracilis. Treponema rigidmn Zinsser and Hop- kins. (Jour. Bact., /, 1916, 489.) From the tissues in five different strains of rab- bit syphilis. Probably a synonym of Treponema cuniculi. Treponema spermiformis Duboscq and Grasse. (Arch. Zool. Exper et G^n., 66, 1927,483.) From the rectum of a termite, Glyptotermes iridipennis. Treponema squatarolae Lebailly. (Compt. rend. Soc. Biol., Paris, 75, 1913, 389.) From the caecum of a bird, Squala- rola squafarola. Treponema stylopygae Dobell. (Do- bell, Arch. f. Protistenk., 26, 1912, 117; Spirochaeta stylopygae Zuelzer, 1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1685.) From the intestines of the cockroach, Stylopyga orientalis. Treponema tricalle Cohn. (Cohn, 1872, quoted from Castellani and Chal- mers, Man. Trop. Med., 2nd ed., 1913, 414.) Treponema triglae Duboscq and Le- bailly. (Arch. Zool. Exp^r et G^n., 10, 1912,331.) From the rectum of a fish, Trigla lucerna. Treponema tropiduri Neiva, Marques da Cunha and Travassus. (Mem. do Inst. Oswaldo Cruz. 6, 1914, 180.) From the blood of a South American lizard, Tropidurus torquatus. The following species are listed in the index of Castellani and Chalmers, Manual of Tropical Medicine, 2nd ed., 1913, 1718- 1719, but are not mentioned in the text (pp. 136-141) : Treponema bovidae, T. camelidae, T. canidae, T.Jelidae, T. hip- popotami, T. reptilia, T. rhinoceri, T. selachii, T . suidae, T. vngulata and T. ursidae. Genvs III. Leptospira Xoguchi. (Jour. Exp. Med., 25, 1917, 753.) Finely coiled organisms 6 to 20 microns in length. Spirals 0.3 micron in depth and 0.4 to 0.5 micron in amplitude. In liquid medium one or both ends are bent into a semicircular hook each involving iV to i of the organism. Spinning movements in liquid and vermiform in semisolid agar, forward or backward. Seen in living prepara- tions only with dark field. Stain with difficulty except with Giemsa's stain and silver impregnation. Require oxygen for growth. The type species is Leptospira icterohaemorrhagiae (Inada and Ido) Noguchi. 1. Leptospira icterohaemorrhagiae (Inada and Ido) Noguchi. {Spirochaeta icterohaemorrhagiae Inada and Ido, Tokyo Ijishinski, 1915; Inada, Ido, Hoki, Ka- neko and Ito, Jour. Exp. Med., 23, 1916, 377; Spirochaeta icterogenes Uhlenhuth and Fromme, Med. Klin., //, 1915, 1202; Spirochaeta nodosa Huebner and Reiter, Deutsch. med. Wochnschr., 41, 1915, 1275; Noguchi, Jour. Exp. Med., 25, 1917, 755; Spiroschaudinnia icterohaemor- rhagiae Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 447; Trepo- nema icterogenes Gonder and Gross, Arch. f. Protistenk., 29, 1919, 62; Spirochaete ictero-haemorrhagica (sic) Lehmann and Neumann, Bakt. Diag., 6 Aufl., 2, 1920, 810; Treponema ictero-hemorragiae (sic) Brumpt, Nouveau Traite de M(5decine, Paris, 4, 1922, 501 ; Treponema nodosum FAMILY TREPOXEMATACEAE 1077 Brumpt, ibid., 508; Leptospira iclerogenes Ford, Textb. of Bact., 1927, 994; Lepto- spira nodosa Ford, ibid., 993.) From Greek icterus, jaundice and hemorrhagiae, bleeding. Morphology : 0.25 to 0.3 by 6 to 9 mi- crons and occasionally 20 to 25 microns. Spiral amplitude: 0.4 to 0.5 micron, regular, rigid. Spiral depth: 0.3 micron, regular. Waves : One or more gentle waves throughout entire length. When in liquid media, one or both ends may be semicircularly hooked, while in semisolid media the organism appears serpentine, waved or bent. Very active fle.xibilitj'. Terminal filament and fiagella absent. Body stains reddish by Giemsa's stain. Bile salts (10 per cent): Easily dis- .solved. Saponin (10 per cent) : Completely re- sistant. Cultured easily in medium containing 10 per cent rabbit serum, 0.2 per cent agar, slight amount of hemoglobin in salt or Ringer's solution. Does not grow in surface colonies. Temperature range: 25° to 37°C. Re- mains alive longer at 25°C. Pathogenic for guinea pigs and deer- mice . Habitat : The cause of infectious jaun- dice in man (Weil's disease). Found in the kidneys, urine and blood of wild rats. No insect vector known. Found free- living in water and slime (in mines). 2. Leptospira hebdomadis (Ido et al.) Xoguchi. (Spirochaeta nanukayami Ido, Hoki, Ito and Wani, Nippon Gakkai Zasshi, 5, 1917, No. 5; Spirochaeta hebdo- madis Ido, Ito and Wani, Jour. Exp. Med., 28, 1918, 435; Spiroschaudinnia hebdomadis Castellani and Chalmers, Man. Trop. Med., 3rd ed., 1919, 448; No- guchi. Jour. Exp. Med., 30, 1919, 17; Treponema hebdomadis Brumpt, Nouveau Traitd de Medecine, Paris, 4, 1922, 501.) From Latin, seven days. Morphologically indistinguishable from Leptospira icteroheamorrhagiae but can be distinguished serologically. In man causes less jaundice than Lepto- spira icterohaemorrhagiae and is never fatal . Identical with Type B, Leptospira au- tumnalis. Slightly pathogenic for young guinea pigs. Is carried by the field vole {Microtus montibelli) . Habitat : Cause of seven-daj^ fever or gikiyami in Japan. 3. Leptospira biflexa (Wolbach and Binger) Noguchi. (Spirochaeta biflexa Wolbach and Binger, Jour. Med. Res., 30, 1914, 23; Xoguchi, Jour. Exp. Med., 27, 1918. 585; Spirochaeta pseiido-icterogenes (aquatilis) Fhlenhuth and Zuelzer, Cent, f. Bakt., I Abt., Orig., 85, 1921,* 141 ; Spi- rochaeta pseudoicterogenes Uhlenhuth and Zuelzer, Klin. Wochmschr., 1, 1922, 2124; Spirochaeta pseudo-icterohemorrhagiae Vinzent, Compt. rend. Soc. Biol., Paris, 05, 1926, 1472; Leptospira pseudoictero- genes Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 461.) From Latin, doubly bent. Size : 0.2 to 0.25 by 5 to 7 microns with tapering ends. Spiral amplitude 0.2 to 0.25 micron. Will pass through an L5 candle filter. Waves : 22 to 30 in number. Stains : Best results with Giemsa's stain. Culture : Can grow in distilled water plus 0.1 per cent potassium nitrate. Rabbit serum in distilled water is best medium. Optimum temperature 20°C. Antigenically distinct from Leptospira icterohaemorrhagiae . Not pathogenic. Source : From tap water, ponds and pools in Berlin. Habitat : Fresh water. 4. Leptospira canicola Okell et al. (Okell, Bailing and Pugh, Vet. Jour., 81, 1925, 3.) From Latin, dog-dweller. 1078 MANUAL OF DETERMINATIVE BACTERIOLOGY Morphologically indistinguishable from Leptospira icterohaernorrhagiae. Cultivation : Same as Leptospira ictero- haernorrhagiae. Immunology : Some cross-reaction with Leptospira icterohaernorrhagiae, but spe- cific in higher dilutions of immune serum. Source : From blood of dogs. Habitat: A natural parasite of dogs. Causes a chronic disease of old dogs char- acterized by uremia, not jaundice. Fatal in 80 per cent of those infected. No intermediate host known. Probably transmitted by direct contact; possibly by healthy carriers. Appendix: The species listed below are inadequately described and may be iden- tical with those described in full. Leptospira aqueductum (sic) Ford. {Spirochaeta pseudoicterogenes aquaeduc- tuum Uhlenhuth and Zuelzer, Cent. f. Bakt., I Abt., Orig., 85, 1921, *150; Ford, Textb. of Bact., 1927, 998.) From fresh water of aqueducts. Probably a synonym of Leptospira biflexa. Leptospira asthenoalgiae Carbo-Noboa. (Bull. Inst. Past., ^;?, 1924, 898.) From blood, urine and organs of persons having dengue. Leptospira autumnalis Topley and Wil- son. (Akiyami Type A, Koshina, Shiwo- zawaand Kitayama, Japan. Med. Wld., 4, 1924, 268; see also Jour. Exp. Med., ^2, 1925, 873; Topley and Wilson, Princip. Bact. and Immun., 1st ed., 2, 1931, 1202; Spirochaeta autumnalis A, quoted from Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 312; Spirochaeta autumnalis Hin- dle, ibid.) The cause of akiyami or har- vest sickness in Japan. May be identical with Leptospira icterohaernorrhagiae. Leptospira bataviae. (1925, quoted from Gispen and Schiiffner, Cent. f. Bakt., I Abt., Orig., lU, 1939, 427.) From a case of fever in the Dutch East Indies. Probably a synonym of Lepto- spira hebdomadis. Leptospira biliohemoglobinuriae (Blan- chard and Lefrou) Noguchi. {Spiro- chaeta bilio-hemoglobinuriae Blanchard and Lefrou, Compt. rend. Acad. Sci., Paris, 175, 1922, 002; Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928,490.) From cases of black- water fever. Leptospira bonariensis Savino and Rennella. (Rev. Inst. Bact. "Dr. Car- los G. Malbram", 12, 1944, 182.) From gray rats. Leptospira bovis Noguchi. (New York State Med. Jour., 22, 1922, 426.) From the gastric mucosa of the ox. Leptospira couvyi Gomes de Faria. (Compt. rend. Soc. Biol., Paris, 90, 1924, 55; Spirochaeta couvyi Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 317.) From the blood of persons having dengue. Leptospira dentate Perrin. (Rev. Mex. de Biol., 2, 1922, 171.) Found in the pus of bucco-maxillary gangrene. Leptospira grippo-typhosa Topley and Wilson. (Topley and Wilson, Princip. Bact. and Immun., 2nd ed., 1936, 728; Spirochaeta dmitrovi Rimpau, Schloss- berger and Kathe, Cent, f . Bakt., I Abt., Orig., HI, 1938, 320.) The cause of swamp fever in Europe. Probably syn- onymous with Leptospira hebdomadis. Also see Baschenin, Cent. f. Bakt., I Abt., Orig., 113, 1929, 438 and 450; Dinger and Verschaffelt, Ann. Inst. Past., ^5, 1930, 396. Leptospira haemoglobinuriae Schiiffner. (Geneesk. Tijdschr. Ned. Indie, 58, 1918, 352; Spirochaeta haemoglobinuriae Hin- dle, Med. Res. Council Syst. of Bact., 8, 1931 , 314. ) From the blood of a Javanese patient suffering from an attack of black- water fever. Leptospira icterohemoglobinuriae Schiiffner. (Schiiffner, Geneesk. Tijd- schr. V. Ned. Indie, 58, 1918, 352, accord- ing to Pettit, Contribution k I'Etude des Spirochetid^s, Vanves, II, 1928; Spiro- chaeta icterohemoglobinuriae Schiiffner, Mededeel. Burgerl. Geneesk. Dienst in Nederl. Indie, 58, 1918, 7 (according to Blanchard and Lefrou, Compt. i-end. Acad. Sci., Paris, 175, 1922, 602) ; Trepo- nema icterohemoglobinuriae Brumpt, FAMILY TREPONEMATACEAE 1079 Nouveau Traits de M^decine, Paris, 4, 1922, 501.) From the blood in a case of blackwater fever. Leptospira interrogans (Stimson) No- guchi. {Spirochaeta interrogans Stim- son, U. S. Public Health Rept., Part 1, 22, 1907, 541; Leptospira icteroides Noguchi, Jour. Exp. Med., 29, 1919, 581; Trepo- nema interrogans Brumpt, Nouveau Traits de Medecine, Paris, 4, 1922, 505; Treponema icteroides Brumpt, ibid. ; Spi- rochaeta icteroides Lehmann and Neu- mann, Bakt. Diag., 7 Aufl., ^,1927, 576; Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Immun., 1928, 454.) Noguchi (1928) regards this species as identical with Leptospira icterohaemor- rhagiae. Leptospira pettiti (Fiessinger) Hindle. (Spirochaete pettiti Fiessinger, Ann. de M^d., 5, 1918, 156; Treponema pettiti Brumpt, Nouveau Traits de Medecine, Paris, 4, 1922, 510; Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 316 ; Spiro- chaeta pettiti Hindle, idem; not Spiro- chaeta pettiti Row, Jour. Trop. Med. and Hyg., 1922, 364.) From urine. Morpho- logically indistinguishable from Lepto- spira icterohaemorrhagiae . Leptospira pyrogenes Vervoort. (Ver- voort, Geneesk. Tijdschr. v. Ned. Indie, 63, 1923, 800; Spirochaeta febrilis Ver- voort, Rep. Far East. Assoc. Trop. Med., London, 1923, 683; Spirochaeta pijrogencs Hindle, Med. Res. Council Syst. of Bact., 8, 1931, 314.) From the blood of persons suffering from dengue-like fevers in Su- matra. Pathogenic. Leptospira saxkoebing Petersen. (Acta Path, et Microbiol. Scand., 21, 1944, 165.) A new serological type. Leptospira salina Ford. (Spirochaeta pseudoicterogenes salina Uhlenhuth and Zuelzer, Cent. f. Bakt., I Abt., Orig., 85, 1921, *150; Ford, Textb. of Bact., 1927, 998.) From salt water. Leptospira trimerodonta (Hoffmann) Noguchi. {Spirochaeta trimerodonta Hoffmann, Deutsch. med. Wochnschr., 46, 1920, 257; Leptospira dentium Hoff- mann, ibid., 625; Leptospira buccalis Fontana, according to Pettit, Contribu- tion a I'Etude des Spiroch^tid^s, Vanves, II, 1928, 232; Noguchi, in Jordan and Falk, Newer Knowledge Bact. and Im- mun., 1928, 461.) From the oral cavity. May be synonymous with Leptospira icterohaemorrhagiae. Spirochaeta anthropopitheci Wilbert and Delorme. (Ann. Inst. Past., 41 > 1927, 1147.) Pathogenic for chimpanzees in French Guinea. Probably identical with Leptospira icterohaemorrhagiae . Spirochaeta elusa Wolbach and Binger. (Wolbach and Binger, Jour. Med. Res., 30, 1914, 9; Treponema elusum Bergey et al.. Manual, 1st ed., 1923, 428.) From pond water. Not pathogenic. For a de- scription of this species, see Bergey et al., Manual, 5th ed., 1939,957. Spirochaeta ictero -uraemia canis Klar- enbeek. (Tijdschr. Diergeneesk., 55, 1928, 227.) From the kidneys of dogs. Pathogenic for guinea pigs. May be syn- onymous with Leptospira icterohaemor- rhagiae or L. canicola. Spirochaeta psendohebdomadis Zuelzer. (1925, in Prowazek, Handb. d. path. Protoz., 3, 1931, 1671.) Probably identi- cal with Leptospira hebdomadis. Spirochaeta trimeres Hoffmann. (Deutsch. med. Wochnschr., 46, 1920, 257.) From the oral cavity. May be synonymous with Leptospira trimero- donta. SUPPLEMENT NO. 1 ORDER RICKETTSIALES April, 1947. FAMILY RICKETTSIACEAE 1083 ORDER RICKETTSIALES GIESZCZYKIEWICZ. (Bull. Intern. Acad. Polon. Sci., Classe Math. Nat., B(l), 1939, 9-30.) Small, rod-shaped, coccoid, spherical and irregularly-shaped microorganisms which stain lightly with aniline dyes. Gram-negative. Usually not filterable. Cultivated outside the body, if at all, only in living tissue, embryonated eggs or rarely in media containing body fluids. Parasitic organisms intimately associated with tissue cells and erythrocytes, chiefly in vertebrates and often in arthropods which act as vectors. The intracellular parasites of Protozoa may also belong here. May cause diseases in man or animals, or both. Key to the families of order Rickettsiales. I. Intracellular parasites, or parasites intimately associated with tissue cells. Do not occur in erythrocytes. Frequently cause diseases of vertebrates trans- mitted by arthropod vectors. Family I. Rickettsiaceae, p. 1083. II. Facultative intracellular or extracellular parasites found characteristically in or on the erythrocytes of vertebrates. May be transmitted by arthropod vectors. Family II. Bartonellaceae, p. 1100. III. Intracellular parasites found in vertebrate tissues and not transmitted by arthropod vectors. Family III. Chlamydozoaceae ,p . llli. *PAMILY I. RICKETTSIACEAE PINKERTON. (Pinkerton, Parasitology, 28, 1936, 186; Rickettsiales Buchanan and Buchanan, Bacteriology, 4th ed., New York, 1938, 49.) Small, often pleomorphic, rod-shaped, ovoid, coccoid and coccus-shaped bacterium- like organisms, intimately associated with arthropod tissues, usually in an intra- cellular position. Stain lightly with aniline dyes. Gram-negative. Have not been cultivated to date in cell-free media. May be parasitic to man and other animals causing diseases (typhus and related ills) that are transmitted by arthropod vectors (lice, fleas, ticks, mites and probably other ectoparasites). Key to the genera of family Rickettsiaceae.f I. Cells rod-shaped, ellipsoidal and coccoid. A. Non-filterable. B. Filterable. II. Cells spherical, occasionally elongated. Genus I. Rickettsia, p. 1084. Genus II. Coxiella, p. 1092. Genus III. Cowdria, p. 1094. * Prepared by Dr. Ida A. Bengtson (retired), National Institute of Health, Bethesda, Mar3'land, November, 1946. Through the courtesy of Dr. Edward A. Steinhaus much use was made of material from his book, Insect Microbiology, Ithaca, 1946, 763 pp. before it was generally available. t Includes only those rickettsiae which have been rather completely studied. For additional rickettsiae, see appendix. 1084 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus I. Rickettsia da Rocha-Lima. (Berl. klin. Wchnschr., 53, 1916, 567-569.) Named for Howard Taylor Ricketts who lost his life studying typhus fever. Small, often pleomorphic, rod-shaped to coccoid organisms occurring intracyto- plasmically in lice, fleas, ticks and mites, or sometimes intranuclearly. Stain lightly with aniline dyes. Gram-negative. Non-filterable. Have not been cultivated in cell-free media. Parasites of man and animals which are the etiological agents of epidemic typhus, murine or endemic typhus, Rocky Mountain spotted fever, tsu- tsugamushi disease, rickettsialpox and other diseases. For reasons that are discussed elsewhere (Bengtson, Jour. Bact., 53, 1947, 325) the genus Dermocentroxenus has been united with the genus Rickettsia. The type species is Rickettsia proivazekii da Rocha-Lima.* Key to the species of genus Rickettsia. I. Louse-borne. II. Flea-borne. III. Tick-borne. IV. Mite-borne. 1. Rickettsia prowazekii. 2. Rickettsia typhi. 3. Rickettsia rickettsii. 4. Rickettsia conorii. 5. Rickettsia tsutsugamushi . 6. Rickettsia akari. 1. Rickettsia prowazekii da Rocha- Lima. (da Rocha-Lima, Berl. klin. Wchnschr., 53, 1916, 567; Rickettsia exanthematotyphi Kodama, Kitasato Arch. Exper. Med., 9, 1932, 360; Rickett- sia prowazeki var. prowazeki Pinkerton, Parasitology, 28, 1936, 186; Rickettsia prowazeki sub-species prowazeki Philip, Amer. Jour. Hyg., 37, 1943, 307.) Named for S. von Prowazek who lost his life studying typhus fever. Minute coccoid, ellipsoidal and ovoid forms to short rods, sometimes long rods and occasionally filamentous forms, often in pairs and occasionally in chains. In infected lice the minute coccoid and paired coccoid forms predominate over the short and long rods and the filamen- tous forms which are up to 40 microns in length. Single elements 0.25 by 0.4 to 0.3 by 0.45 micron. Pairs range from 0.25 by 0.7 to 0.3 by 1.1 microns. In yolk sacs the organisms vary in size from minute coccoid forms in heavily infected tissue to rod forms resembling small bacteria in lightly infected tissue. Within the same smear of infected mam- malian cells and in chick embryo tissue the organisms are quite uniform in size and morphology. Occur intracytoplas- mically in vascular endothelial cells and in serosal cells. Non-motile. * The Editors of the Manual follow Recommendation XL of the International Botanical Code (see p. 59) in regard to the endings used for specific names. This calls for the use of the ii ending for epithets taken from the name of a man ending in a consonant (except names ending in er). Some students of the Rickettsiaceae fol- low the International Rules of Zoological Nomenclature which use ii only in case the name used was employed and declined in Latin. Zoologists use the single i for modern patronymics based on all other names of men. FAMILY RICKETTSIACEAE 1085 The organisms are colored purplish with the Giemsa stain, the two indi- viduals of a pair being connected by a zone of faintly blue stained material. They are colored blue with Castaiieda stain (Jour. Inf. Dis., 47, 1930, 416) and bright red against a blue background with Machiavello stain (Rev. Chilena de Hig. y Med. Prev., 1, 1937, 101). Gram- negative . Cultivation : In plasma tissue cultures of mammalian cells, in the louse intestine, in modified Maitland media with and without agar, on chorioallantoic mem- brane and yolk sac of chick embryo, the latter being currently the medium of choice. Optimum temperature 32°C in plasma tissue culture, 35°C in chick embryo cells. Immunology : Immunity prolonged but may not be complete in man. Indis- tinguishable from endemic (murine) typhus in cross immunity tests in guinea pigs, but distinguishable from Rocky Mountain spotted fever and other rickett- sial diseases in such tests. Neutraliz- ing antibodies are found in the serum of recovered guinea pigs and convalescent humans up to 2 to 3 weeks after defer- vescence. Killed vaccines produced from infected lice and from infected yolk sacs afford a high degree of protection against the disease. Hyperimmune anti- sera for therapeutic use have been pro- duced in rabbits by injection with infected yolk sac suspensions and in horses and donkeys with infected mouse lung suspensions. Serology : Strains from various parts of the world are closely related as de- termined by complement fixation; are distinguishable from other rickettsiae by agglutination, complement fixation and precipitin tests; have a common anti- genic factor (alkali stable polysac- charide) with Proteus 0X19; and have a soluble antigen in yolk culture. Lethal effect : Heavily infected j^olk sac cultures injected intravenously or intraperitoneally are fatal to white mice in a few hours. Resistance to chemical and physical agents : Readily inactivated by heat and chemical agents. A temperature of 50°C kills the organism in 15 to 30 minutes, and 0.5 per cent phenol and 0.1 per cent formalin kill the organism. Pathogenicity : Pathogenic for man, apes, monkej's, guinea pigs, cotton rats, gerbilles, the louse (Pediculus humanus). Inapparent infections occur in white mice, white rats and rabbits. A charac- teristic febrile reaction with no mortality and without testicular swelling occurs in the guinea pig. Passage in guinea pigs is accomplished by transfer of blood or brain from infected animals. Causes a febrile disease with e.xanthema and high mortality in man. Source : Seen in the blood of tj'phus patients and in smears of epithelial cells of the intestinal tract of lice fed on typhus patients. Habitat : The body louse (Pediculus humanus var. corporis), head louse {Pediculus humanus var. capitis) and Pedicinus longiceps. The etiological agent of epidemic typhus (European typhus, classical typhus, typhus exan- thematicus). 2. *Rickettsia typhi (Wolbach and Todd) Philip. {Dermacentroxenus * Some may regard the binomial Rickettsia typhi as invalid because of its previous use by do Amaral and Monteiro for the organism causing eastern Rocky Mountain spotted fever. However, because the binomial Dermacentroxenus typhi Wolbach and Todd clearly has priority and because the binomial proposed by do Amaral and Monteiro has never come into general use, Rickettsia typhi Philip has been accepted for use in the Manual. If Philip's binomial had been rejected, then it would have been necessary to accept Rickettsia nianchuriae Kodama et al. as this appears to have priority over the more generally used Rickettsia mooseri Monteiro. — Editors. 1086 MANUAL OF DETERMINATIVE BACTERIOLOGY typhi Wolbach and Todd (not Tood), Ann. Inst. Past., 34, 1920, 158; minute intracellular bodies, Mooser, Jour. Inf. Dis., 43, 1928, 261 ; Rickettsia manchuriae Kodama, Takahashi and Kono, Saiking- aku-Zasshi (Jap.), No. 426, 427, Aug. and Sept., 1931; see Kodama, Kono and Takahashi bibliography, Kitasato Arch. Exper. Med., 9, 1932, 95; Rickettsia mooseri Monteiro, Mem. Inst. Butantan, 6, 1931, 97 (pub. July, 1932), see do Amaral and Monteiro, bibliography, ibid., 7, 1932, 367; Rickettsia exanthematofebri Kodama, Kitasato Arch. Exp. Med., 9, 1932, 360; Rickettsia muricola Monteiro and Fonseca, Brazil Med., 46, 1932, 1032; Rickettsia murina and Rickettsia fletch- eri Megaw, Trans. Roy. Soc. Trop. Med. Hyg., 29, 1935, 105; Rickettsia prowazeki var. mooseri Pinkerton, Para- sitology, £8, 1936, 185; Rickettsia prowa- zeki sub-species typhi Philip, Amer. Jour. Hyg., 37, 1943, 304; Rickettsia typhi Philip, idem; not Rickettsia typhi do Amaral and Monteiro, Rev. Sud. Am^r. de M^d. et Chirug., 4, 1933, 806.) From M. L. typhus, typhus. Resembles Rickettsia prowazekii in morphological and staining properties. Non-motile. Gram-negative. Cultivation : May be cultivated in plasma tissue culture of mammalian cells, in modified Maitland media with and without agar, in fleas, in the peri- toneal cavity of X-rayed rats, in the lungs of white mice and in white rats following intranasal inoculation, in the lungs of rabbits following intratracheal inoculation, in the chorio-allantoic mem- brane and the yolk sac of the chick embryo. Optimum temperature 35°C in chick embryo cells. Immunology : Prolonged immunity in man and animals following infection. Complete cross immunity between epi- demic and endemic typhus in guinea pigs recovered from infections with Rickettsia protmzekii and Rickettsia typhi. No cross immunity between en- demic typhus and Rocky Mountain spotted fever, Q fever or tsutsugamushi disease in guinea pigs. Serology: Distinguishable from the rickettsiae of spotted fever, Q fever and tsutsugamushi disease by complement fixation, agglutination and precipitin tests, less readily from R. prowazekii by these tests. Has common antigenic fac- tor with Proteus 0X19, and soluble antigen in yolk-sac cultures. Lethal effect: Heavily infected yolk sac cultures injected intravenously or intraperitoneally fatal to white mice in a few hours. Pathogenicity : Pathogenic for man, apes, monkeys, rabbits, guinea pigs, white rats, eastern cotton rat, white mice, gerbilles. Other susceptible ani- mals include the woodchuck, house mouse, meadow mouse, white-footed mouse, old-field mouse, cotton mouse, golden mouse, wild rat (Rattus norvegi- cus), wood rat, rice rat, flying squirrel, gray squirrel, fox squirrel, gophers, cot- ton-tail rabbit, swamp rabbit, chipmunk, skunk, opossum and cat. A characteris- tic febrile reaction occurs in the guinea pig with testicular swelling without ulceration, after intraperitoneal inocula- tion. Passage in guinea pigs is accom- plished by transfer of testicular wash- ings or blood from infected animals. Cause of a febrile disease with exanthema in man, with low mortality. Source : Seen by Wolbach and Todd {loc. cit.) in the endothelial cells of the capillaries, arterioles and veins in sec- tions of skin from cases of Mexican typhus (tabardillo). Also described by Mooser {loc. cit.) in sections and smears of the proliferated tunica vaginalis of guinea pigs reacting to the virus of Mexi- can typhus. Habitat: Infected rat fleas {Xeno- psylla cheopis, Xenopsylla astia), in- fected chicken fleas {Echidnophaga galli- nacea) found on wild rats, and the rat louse {Polyplax spinulosiis) . Wild rats and field mice act as the reservoir of infection. The etiological agent of en- FAMILY RICKETTSIACEAE 1087 demic (murine) typhus which is trans- mitted to man by the rat flea. 3. Rickettsia rickettsii (Wolbach) Brumpt. {Dermacentroxcnus rickettsi Wolbach, Jour. Med. Res., U, 1919-20, 87; Rickettsia rickettsi Brumpt, Precis de Parasitologie, 3rd ed., 1922, 757; Rickettsia brasiliensis Monteiro, Mem. Inst. Butantan, 6, 1931, 3; * Rickettsia typhi do Amaraland Monteiro, Rev. Sud. Am^r. de M6d. et Chirurg., 4, 1933, 806; Dermacentroxenus rickettsi var. brasili- ensis Pinkerton, Parasitology, 28, 1936, 186.) Rickettsia dermacentroxenus, a corruption of Dermacentroxenus rickettsi, though widely used, has no genuine taxonomic standing. Named for Howard Taylor Ricketts, who first transmitted the disease from human cases to monkeys and guinea pigs with the production of characteristic symptoms and lesions and fatal effect. Minute paired organisms surrounded by a narrow clear zone or halo and often lanceolate, resembling in appearance a minute pair of pneumococci. Approxi- mately 0.2 to 0.3 micron by 1 micron. Non-motile. In smears of mammalian tissues there occur in addition to the lanceolate forms, slender rod-shaped forms stained blue with the Giemsa stain, sometimes ex- hibiting polar granules, stained purplish or reddish. There are also minute pale blue-staining rounded forms. In the tick there are three forms: (1) Pale blue bacillary forms curved and club-shaped, (2) smaller bluish rods with deeply staining chromatoid granules and (3) more deeply staining, purplish, lanceo- late forms. A very minute form may appear in tightly packed masses in the nuclei of the cells. Occurs in the cyto- plasm and nucleus in all types of tissue in the tick and in the vascular endothe- lium, in the serosal cells of the peritoneal cavity, in the smooth muscle cells of arteriolar walls and in the macrophages of mammals. In yolk sac cultures and in the Mait- land media cultures, bacillary forms often occur in pairs. In single smears from infected yolk sacs, the rickettsiae are rather uniform in size and morphology and are definitely lai'ger than Rickettsia prowazekii and Rickettsia typhi. They also grow more sparsely. Stain blue with the Castaiieda stain and bright red against a blue background of tissue with the Machiavello stain. Cultivation : May be cultivated in plasma tissue culture of mammalian cells, in Maitland media with and with- out agar, on the chorio-allantoic mem- brane and in the yolk sac of the chick embryo, and in ticks. Optimum temperature 32°C in plasma tissue culture, 35°C in chick embryo cells. Immunology : Prolonged immunity in man and animals after recovery from in- fection. Killed vaccines produced from infected ticks and from infected yolk sacs afford considerable protection against the disease. Therapeutic anti- sera have been produced by the injection of rabbits with tick virus and with in- fected yolk sac. No cross immunity between spotted fever in guinea pigs recovered from infections with Rickett- sia rickettsii and typhus in guinea pigs recovered from infections with Rickettsia prowazekii and Rickettsia typhi. Cross immunity between spotted fever in guinea pigs recovered from infections with Rickettsia rickettsii and bouton- neuse fever in guinea pigs recovered from infections with Rickettsia conorii, but spotted fever vaccine does not protect against boutonneuse fever of the Medi- terranean area or against infections with the South African strains of Rickettsia conorii. * Erroneously applied by do Amaral and Monteiro to the so-called eastern type of Rocky Mountain spotted fever. — Editors. 1088 MANUAL OF DETERMINATIVE BACTEEIOLOGY Serology : Distinguishable from Rick- ettsia prowazekii and Rickettsia typhi by complement fixation and agglutination with specific antigens. Distinguishable from Rickettsia conorii by complement fixation, though some degree of cross fixation indicates antigenic relationship. Has common antigenic factor with Pro- teus 0X19 but not distinguishable from Rickettsia prowazekii and Rickettsia typhi by Weil-Felix test. Resistance to chemical and physical agents : Readily inactivated by heat and chemical agents. Destroyed by a tem- perature of 50°C in 10 minutes, and by 0.5 per cent phenol and 0.1 per cent formalin. Destroyed by desiccation in about 10 hours. Pathogenicity : Pathogenic for man, monkeys and guinea pigs. Rabbits and white rats are moderately susceptible. Animals susceptible in varying degree.s include species of ground squirrels, tree squirrels, chipmunks, cotton-tail rabbits, jack rabbits, snowshoe rabbits, marmots, wood rats, weasels, meadow mice and deer mice. In Brazil the opossum, rab- bit, dog and cavy have been found natu- rally infected and the Brazilian plains dog, capybara, coati and certain bats are also susceptible. Sheep are mildly susceptible. A febrile reaction occurs in guinea pigs with typical scrotal lesions, involving petechial hemorrhages in the skin, which may become necrotic. Virulent strains kill 80 to 90 per cent of the animals, milder strains kill 20 to 25 per cent. Passage in guinea pigs is accomplished by transfer of blood from infected animals. A febrile reaction accompanied by exan- thema occurs in man. Mortality is high in some localities, low in others. Source : Seen by Ricketts (Jour. Amer. Med. Assoc, 52, 1909, 379) in the blood of guinea pigs and monkeys experimen- tally infected with Rocky Mountain spotted fever and in the salivary glands, alimentary sac and ovaries of infected female Dcrmacentor ticks and in their ova. Habitat: Infected wood tick {Derma- centor andersoni) and the dog tick {Der- macentor variabilis), also the rabbit tick (Haemaphysalis leporis-palustris) , Am- blyonima hrasiliensis, Amhlijomma cajennense, Amblyonwia striatum, Am- blyomma americanuni and Ixodes denta- tus. A number of ticks belonging to the genera Amblyomma, Dermacentor, Rhipi- cephalus, Ornithodoros and Haema- physalis have been experimentally infected. The virus is transmissible through the ova of female ticks. The etiological agent of Rocky Mountain spotted fever, Sao Paulo exanthematic typhus of Brazil, Tobia fever of Colombia and spotted fever of Minas Geraes which are all transmitted to man by the bite of infected ticks. i. Rickettsia conorii Brumpt. (Brumpt, Compt. rend. Soc. Biol., Paris, 110, 1932, 1199; Rickettsia megawi var. pijperi do Amaral and Monteiro, Mem. Inst. Butantan, 7, 1032, 361 ; Rickettsia blanci Caminop^tros, l'"' Cong. Internat. Hyg. Mediterr., Rapports et Compt. rend., 2, 1932, 202; Dermacentroxenus rickettsi var. pijperi Mason and Alexan- der, Onderst. Jour. Vet. Sci. and An. Indust., 13, 1939, 74; Dermacentroxenus rickettsi var. conori Mason and Alex- ander, ibid. ; Dermocentroxenus conori Steinhaus, Insect Microbiology, 1946, 339.) Named for A. Conor who with A. Bruch published in 1910 the first clini- cal description of boutonneuse fever. Resembles Rickettsia rickettsii. In the tick, diplococcoid and diplobacillary forms predominate, though when the rickettsiae occur in compact masses they are smaller and more coccoid. In tissue cultures the organisms are lanceolate, diplococcoid, and diplobacillary, oc- curring in the nuclei as well as in the cytoplasm of the cells. Size 0.3 to 0.4 by 1 to 1.75 microns. Non-motile. Stain purplish with the Giemsa stain, FAMILY RICKETTSIACEAE 1089 blue with the Castaiieda stain and bright red with a blue background with the Machiavello stain. Gram-negative. Cultivation : May be cultivated in plasma tissue culture of mammalian cells, in modified JNIaitland media, and in the yolk sacs of chick embryos. Immunology : The disease is related immunologically to Rocky Mountain spotted fever with which it cross im- munizes, but the spotted fever vaccine does not protect against the Mediterra- nean and South African strains of bou- tonneuse fever. Serology : Distinguishable from Rick- ettsia rickettsii by complement fixation. Has a common antigenic factor with Proteus 0X19 and 0X2. Pathogenicity : Pathogenic for man and guinea pigs. It is also pathogenic in varying degrees for dogs, horses, spermo- philes, monkeys, rabbits, gerbilles and white mice. Boutonneuse fever is a much less virulent infection for the guinea pig than Rocky Mountain spotted fever. A tem- perature reaction occurs, accompanied by scrotal swelling but there is no slough- ing. There is practicallj^ no mortality. Passage in guinea pigs is accomplished by transfer of blood from an infected animal . In man, localized primary sores (taches noires) and an inflammatory reaction in the regional lymph nodes occur at the site of the tick bite. A febrile reaction with exanthema occurs and mortality is low. Source : Seen by Caminop^tros (Compt. rend. Soc. Biol., Paris, 110, 1932, 344) in smears from the tunica vaginalis of guinea pigs inoculated with infected dog ticks {Rhipiccphalus sanguineus) . Habitat: The brown dog tick (Rhipi- cephalus sanguineus) and also the ticks, Amhlyomma hebraeum, Haemaphysalis leachi, Rhipicephalus appendiculatus and Boophilus decolor atus. Transmissible through the ova of adult female ticks. The probable animal reservoir is the dog. The etiological agent of boutonneuse fever in man, also known as eruptive, Mediterranean or Marseilles fever and probably Kenya typhus and South Afri- can tick bite fever, though the identity of the latter with boutonneuse fever has been questioned. 5. "Rickettsia tsutsugamushi (Haya- shi) Ogata. (Theileria tsutsugamushi ° Some may question the use of this binomial on the ground that Hayashi thought that this species was possibly or probably protozoan in nature when he proposed the name Theileria tsutsugamushi (loc. cit.) in 1920. However he questions whether Theileria is the correct generic name in this paper and accepts the viewpoint that this organism is a rickettsia in a paper published in 1924 entitled. On Rickettsia, Trans. Jap. Path. Soc, 14, 1924, 198-201. He does not use the binomial Rickettsia tsuts^iga- mushi in this paper as indicated by some of his friends in latter papers (Ogata, loc. cit., Ivawamura, loc. cit.) and apparently first uses it himself in a paper entitled, On Tsutsugamushi Disease, Jap. Path. Soc, 22, 1932, 686. Hayashi was not the first to recognize the probable rickettsial nature of the organ- ism of the tsutsugamushi disease (see Blake et al., Amer. Jour. Hyg., 41, 1945, 257-262) and some even question whether any of the bodies that he found in human lympho- cytes from lymph nodes, in mononuclear endothelial phagocytes of the spleen and lymph nodes, and in tissues taken from 1 he region of the mite bite in patients suffering from tsutsugamushi fever were the same as organisms described as Rickettsia orien- talis by X'agayo et al. (loc. cit.). This position is not supported, however, by Nagayo and his associates who admit that their organisms are identical with some of the organisms described by Hayashi. Mitamura (Trans. Jap. Path. Soc, 21. 1931, 463) sums this up as follows: "Wir stellen 1090 MANUAL OF DETERMINATIVE BACTERIOLOGY Hayashi, Jour. Parasit., 7, 1920, 63; troxenus orientalis Moshkovsky,Vspekhi *Rickettsia orientalis Nagayo, Tamiya, Souremennoi Biologii (Russian) (Ad- Mitamura and Sato, Jikken Igaku vances in modern biology), 19, 1945, 13.) Zasshi, ^.^, May 20, 1930,8 pp.: fiJicA-e^^sia From two Japanese ideographs trans- tsutsugamushi Ogata, Cent. f. Bakt., literated tsutsuga, something small and I Aht., 122,1931,249; Rickettsia akamushi dangerous, and mushi, a creature now Kawamuraand Imagawa, ihid., 122, 1931, known to be a mite. If the i ending is 258; Rickettsia orientalis var. schuffneri accepted as forming a Latin genitive, do Amaral and Monteiro, Mem. Inst. the modern meaning of the species name, Butantan, 7, 1932, 360; Rickettsia meg- tsutsugamushi , would be 'of a dangerous awi do Amaral and Monteiro, idem; mite'. Rickettsiamegawivav. fletcheri do Amaval Small pleomorphic bacterium-like mi- and Monteiro, ibid., 361; Rickettsia croorganisms, usually thicker than ^z'cA;- ts utsugamushi -orientalis Kawainursi,lSi is- ettsia proivazekii, Rickettsia typhi, Rick- shin Igaku, 23, 1934, 000; Rickettsia pseu- ettsia rickcltsii and Coxiella burnetii and dotyphi Vervoort, see Donatien and less sharply defined. Ellipsoidal or rod- Lestoquard, Acta Conv. Tertii Trop. shaped, often appearing as a diplococcus atque malariae morbis, pars I, 1938, or as a short bacillus with bipolar stain- 564; Rickettsia sumatranus (sic) Kou- ing resembling the plague bacillus. Dif- wenaar and Wolff, Proc. 6th Pacific Sci. fusely distributed in the cytoplasm of Cong. (1939), 5, 1942, 636; Dermacen- the cell. Size 0.3 to 0.5 by 0.8 to 2 nicht in Abrede dass Herr Hayashi bei einem kleinen Teil der von ihn beschriebenen Korperchen unsere Rickettsia orientalis vor sicli gehabt hat". Hayashi vigorously defends his own observations in the same discussion and the following year after making comparative studies of strains of Rickettsia orientalis and his own Rickettsia tsutsugamushi reaches the following conclusion {loc. cit.) "Rickettsia tsutsugamushi and Rickettsia orientalis refer to one and the same species of microorganisms and there seems to be no way in which one can be recognized as differing from the other." Under these conditions the only valid name appears to be Rickettsia tsutsugamushi. — Editors. * These authors publish practically the same preliminary paper in three other places as follows: Compt. rend. Soc. Biol., Paris, 104, June 14, 1930, 637-641; Jap. Jour. Exper. Med., 8, Aug. 20, 1930, 309-318 and Trans. Jap. Path. Soc, 20, 1930, 556-566. The complete report on this work did not appear until the following year: Jap. Jour. Exper. Med., 9, iMarch 20, 1931, 87-150.— Editors. t This binomial apparently first appears in the literature in a review article by Kawamura (Handbuch der path. Microorganismen, Kolle and Wassermann, 3 Aufl., 8, 1930, 1398) where it is used incidentally and is attributed to Hayashi, 1923. The fact that Hayashi did not use Rickettsia tsutsugamushi before 1931 is confirmed by Mitamura (Trans. Jap. Path. Soc, 21. 1931, 463) who states in a footnote: Kawamura und Ogata geben an, dass Hayashi 1923 fiir den Erreger den Namen Rickettsia tsutsu- gamushi vorgeschlagen hat. Eine solche Angabe Hayashi, is nicht nur uns, sondern auch dem Autor, wie er uns personlich erzahlt, unbekannt." Ogata apparently first used Rickettsia lsufsuga7nushi in the title of a paper that he presented in 1930 to the 8th Cong. Far East Assoc. Trop. Med. which, however, appeared in the Transactions of the Congress, 2, June, 1932, 167-171. Meanwhile, the same paper with an added discussion of the nomenclature appeared in the Cent. f. Bakt., I Abt., Orig., 122, Oct. 1, 1931, 249-253 and it is this paper that is usually regarded as establishing the use of Rickettsia tsutsugamushi for this species. — Editors. FAMILY RICKETTSIACEAE 1091 microns. Non-motile. Colored purplish with the Giemsa stain, and red against a blue background with the Machiavello stain. Stains well with azur III and methylene blue. Gram-negative. Cultivation : In plasma tissue culture of mammalian cells; on the chorio-allan- toic membrane and in the yolk sac of the chick embryo; in rabbit testes and in the endothelial cells overlying Descemet's membrane of the rabbit eye. Immunology : Immunity conferred b}' infection appears less complete than in typhus and Rocky IMountain spotted fever. Strains from several different areas have been found to cross immunize in guinea pigs, but the true relationship of the disease occurring in different localities remains to be determined. Reciprocal cross-immunity between mite strains and human strains has been demonstrated in rabbits, hamsters and mice. Serology : Antigens from different strains vary in sensitivity when tested bv complement fixation with immune .ciera. There are probably a number of different types on the basis of comple- ment fixation with immune sera. Has a pr^mmon antigenic factor with Proteus OY-K. Re.sistance to chemical and physical aeents : Readily inactivated by heat and chemical agents. Destroyed by a tem- perature of 50°C for 10 minutes, and by 0.1 per cent formalin and 0.5 per cent phenol. Pathogenicity : Pathogenic for man, monkeys, gibbons, guinea pigs, hamsters, rats, voles, mice, gerbilles, rabbits (by intraocular injection) and chick embryo. There is wide variation in the virulence of different strains for laboratory ani- mals, infection being established with great difficulty with some, while others may cause a high mortality. A febrile reaction occurs in guinea pigs. Passage in guinea pigs and mice is ac- complished by inoculation of infected spleen or blood from an infected animal. passage in rabbits by intraocular inocu- lation of blood, lymph node or organ emulsions of infected animals. Ascites, enlarged spleen often with a fibrinous deposit are characteristic. In man an eschar wifh adenopathy de- velops at the site of the mite bite. In scrub typhus the eschar is not present. A febrile reaction with exanthema occurs and mortality is variable. In rabbits infection of Descemet's membrane follows intraocular injection of infected material. Source : Seen by Hayashi in smears and sections of the lesion (eschar) at the site of the mite bite and in smears and sections of the adjacent lymph nodes from cases of the disease; also seen by Nagaj^o et al. (loc. cit.) in the endothelial cells overlying Descemet's membrane in rabbits inoculated intraocularly with in- fectious material. Habitat: The mites {Trombicula aka- mushi, Tromhicula deliensis syn. T. walchi, Tromhicula fletcheri Sund probably several others). Infective through the ova of the adult female. Only the larvae feed on rodents or man. Reservoir hosts are probablj^ certain wild rodents, in- cluding house and field rats, mice and voles and probably some birds. The etiological agent of tsutsugamushi disease and scrub typhus (for numerous other designations of the disease see Farner and Katsampes, U. S. Naval Med. Bull., 43, 1944, 800). Note : Rickettsia nipponica Sellards. (Sellards, Amer. Jour. Trop. Med., 3, 1923, 545; Rickettsoides nipponica da Rocha-Lima, in Kolle and Wasserman, Handb. d. path. Mikroorganismen, 3 Aufl., 8, 1930, 1350.) This problemati- cal organism was thought by its author to be the cause of tsutsugamushi disease. Because it was cultivatible by the meth- ods used by Sellards, it is not now re- garded as identical with Rickettsia tsutsu- gajnushi Ogata. Rickettsoides nipponica is the type species (monotypy) of the 1092 MANUAL OF DETERMINATIVE BACTERIOLOGY genus Rickettsoides da Rocha-Lima {loc. cit.). 6. Rickettsia akari Huebner, Jellison and Pomerantz. (Pub. Health Pi.ept.,ffi, 1946, 1682.) From ^-1 car as, a genus of mites . Minute diplobacilli, occurring intra- cellularly and extracellularly, and bi- polarly stained rods. Resemble typical rickettsiae morphologically. Non-mo- tile. Stain well by Machiavello's method, the organisms appearing bright red against a blue background. Stain poorly with methylene blue. Gram-negative. Occur intracytoplasmically and have been seen intranuclearly in yolk sac cells. Cultivation : In the yolk sac of the chick embryo. No growth on artificial culture media. Immunology : Guinea pigs recovered from rickettsialpox are immune to infec- tion with strains isolated from infected mites. Serology : Antigens prepared from in- fected yolk sacs are highly specific except for cross i-eactions with Rocky Mountain spotted fever antigens. Sera from con- valescent patients fixed complement with the homologous antigen and usually with Rocky Mountain spotted fever antigens though at a lower titer. Does not have a common antigenic factor with Proteus strains except that low titers were ob- tained in a few recovered cases in agglu- tination tests with Proteus 0X19. Pathogenicity : Pathogenic for man with focal initial erythematous lesion and adenopathy, followed by fever and appearance of macular rash. No mortal- ity. Experimental infections have been produced in white mice and guinea pigs by the inoculation of infected blood (ir- regularly), and of infected liver and spleen suspensions, infected brain, in- fected lymph nodes, tunica washings of infected animals and by infected yolk sacs. Sjnnptoms in mice include inac- tivity, accelerated respiration, ruffled fur, with occasional deaths; in guinea pigs, fever and marked scrotal reactions. Infected embryos are killed in 4 to 7 days. It has not been found pathogenic for monkeys, distinguishing it from Rickettsia conorii. It is also probably more pathogenic for white mice than Rickettsia conorii. Source : Blood of a human case of rickettsialpox in New York City. Habitat : Blood of human cases and an ectoparasite of rodents, the mite {Allo- dermaniissus sanguineus Hirst). The etiological agent of human rickettsialpox. Genus II. Coxiella Bengtson, gen. nov. (Subgenus Coxiella Philip, Amer. Jour. Hyg., 37, 1943, 306; generic status recom- mended by Steinhaus, Insect Microbiology, 1946, 263.) Named for Herald R. Cox who first described the organism in guinea pigs inoculated with infected ticks col- lected in Montana. Small, pleomorphic, rod-shaped and coccoid organisms, occurring intracellularly in the cytoplasm and extracellularly in infected ticks. Stain lightly with aniline dyes. Gram-negative. They are filterable. Have not been cultivated in cell-free media. Parasites of man and animals which include the etiological agent of Q fever. The type species is Coxiella burnetii (Derrick) Bengtson. 1. Coxiella burnetii (Derrick) Bengt- son comb. nov. (Rickettsia burneli Der- rick, Med. Jour. Australia, 1, 1939, 14; Rickettsia diaporica Cox, Pub. Health Rep., 54, 1939, 1826; Rickettsia burneti var. americana, Anon., Brit. Med. Jour., 2, lS\\\,r:>'S^; Rickettsia (Coxiella) burneti Philip, Amer. Jour. Hyg., 37, 1943, 306.) Named for F. M. Burnet who discovered the organism in Australia. FAMILY EICKETTSIACEAE 1093 Small bacterium-like, pleomorphic or- ganisms varying in size from coccoid forms to well marked rods. Occur as cytoplasmic micro-colonies with diffuse or compact distribution of the organisms through the cytoplasm. Also seen ex- tracellularly, where they appear as small lanceolate rods, diplobacilli and occa- sionally segmented filamentous forms. Chains of 3 to 6 elements often seen. Quite uniform in size and morphology in infected yolk sacs and in mouse spleen with exceedingly minute forms in heavily infected material. Small lanceolate rods, 0.25 by 0.4 to 0.5 micron, bipolar forms 0.25 by 1.0 micron, diplobacilli 0.25 by 1.5 microns. Non-motile. With Giemsa's stain they appear red- dish-purple, with Machiavello's stain bright red against a blue background. Gram-negative. Cultivation : May be cultivated in plasma tissue cultures, in modified Mait- land media and in the yolk sac of chick embryos. Immunology: There is complete cross immunity between Australian and American strains of Q fever in guinea pigs. Strains from other parts of the world also ci'oss immunize. Serology : American and Australian strains are identical by agglutination and agglutinin absorption. Strains from various countries are serologically related as shown by complement fixation. Q fever is distinguishable from other rickettsial diseases by complement fixa- tion tests. No common antigenic factor with any Proteus strain has been demon- strated. Filterability : The infectious agent of Q fever readily passes Berkefeld N filters which are impermeable to ordinary bac- teria and W filters which are impermeable to typhus and spotted fever rickettsiae. Resistance to chemical and physical agents: Comparatively resistant to heat, drying and chemical agents. Survives at least 109 days in cell -free media with- out loss of titer, resistant to 60°C for 1 hour and to 0.5 per cent formalin and 1 per cent phenol when tested in fertile eggs. Pathogenicity : Pathogenic for man, guinea pig and the white mouse. The monkey, dog, white rat and rabbit are mildly susceptible. Certain bush ani- mals in Australia, particularly the bandi- coot, are susceptible and these animals have been found naturally infected. Other rodents and marsupials are mildly susceptible. Calves have been experi- mentally infected and cows have been found recovered from naturally acquired infections. A febrile reaction occurs in guinea pigs but mortality is low except with heavily infected yolk sac which causes a high mortality. On subcutaneous or intra- dermal inoculation a marked inflamma- tory thickening of the skin occurs at the site of inoculation. On autopsy the spleen is enlarged from 2 to 12 times by weight and is engorged with blood. Transfer in guinea pigs and mice is ac- complished by transfer of infected liver and spleen. A febrile reaction often ac- companied by pneumonitis occurs in man, but mortality is low. Source : First seen in smears from mice inoculated intraperitoneally with infec- tious material by Burnet and Freeman (Med. Jour. Australia, 2, 1937 (2), 281). Habitat : The wood tick {Dermacentor andersoni) and the ticks, Dermacentor occidentalis, Amblyomma americanum, Haemaphysalis leporis-palustris , Ixodes dentatus and Haemaphysalis humerosa. Several other species of ticks have been shown to transmit experimentally the virus of Q fever. It has been found to survive in the ova of the female ticks (Dermacentor andersoni and Haemaphy- salis humerosa) . The bandicoot {Isodon macrurus) is probably the natural reser- voir of the disease in Australia. The etiological agent of Q (Queensland) fever 1094 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus III. Cowdria Bengtson, gen. nov. Named for E. V. Cowdry who first described the organism in heartwater of three ruminants, sheep, goats and cattle. Small pleomorphic, spherical or ellipsoidal, occasionally rod-shaped organisms, occurring intracellularly in ticks. Gram-negative. Have not been cultivated in cell-free media. Parasites which are the etiological agent of heartwater of cattle, sheep and goats. The type species is Cowdria ruminantiwn (Cowdry) Bengtson. 1. Cowdria ruminantiixm (Cowdry) Bengtson, comb. nov. (Rickettsia rumi- naniium Cowdry, Jour. Exp. Med., 4^, 1925, 231; Rickettsia (Coivdria) rurni- nantiu7n Moshkovsky, Uspekhi Soure- mennoi Biologii (Russian) (Advances in Modern Biology), 19, 1945, 18.) From M. L. Ruminantia, the cud-chewing mammals. Differ morphologically from typical rickettsiae, showing usually spherical and ellipsoidal forms; occasionally bacil- lary forms. Irregular pleomorphic forms occur. Grow in the cytoplasm of cells, sometimes in densely packed masses. Size of cocci from 0.2 to 0.5 micron in diameter in the endothelial cells of ani- mals, 0.2 to 0.3 micron in diameter in ticks. Bacillary forms 0.2 to 0.3 by 0.4 to 0.5 micron and pairs 0.2 by 0.8 micron in ticks. Non-motile. Stain blue with the Giemsa stain and can also be stained by methylene blue and other basic aniline dyes. Gram- negative. Cultivation not reported. Immunology : Immunity incomplete after recovery from the infection. The organisms are found in the tissues long after recovery. There is some evidence of a variety of strains. Pathogenicity: Pathogenic for goats, sheep and cattle. Transmissible to goats by inoculation of infected blood intra- jugularly. The most characteristic le- sion is the hydropericardium of infected animals. The only small animal shown to be susceptible is the ferret. Source : Seen in the endothelial cells of renal glomeruli and in the endothelial cells of the cerebral cortex of animals suf- fering from heartwater and in the tick, Amhlyoynma hebraeuni. Habitat : The bont tick {Amhlyomma hebraeiun) and also Amblyomma variega- tum. When the tick is infected in the larval state, it can transmit the infection to the nymphal and adult stages, but the disease is not transmissible through the ova of the adult female tick. The etio- logical agent of heartwater in sheep, goats and cattle in South Africa. Appendix I: Further studies of the organism of trench fever are required be- fore the relationship of Rickettsia quin- iana to the other more firmly established species of rickettsiae can be determined. Therefore, it is placed in this appendix. 1. Rickettsia quintana Schminke. (Schminke, Miinch. med. Wchnschr., 64, July 17, 1917, 961; Rickettsia wolhynica Jungmann and Kuczynski, Ztschr. klin. Med., 85, 1918, 261; Fossilis quintana suggested as a possible subspecies "if necessary" by Megaw, Trop. Dis. Bull., 40, 1943, 828.) Probable synonym : Rickettsia pediculi Munk and da Rocha-Lima, Miinch. med. Wchnschr., ^4, 1917, 1423. Coccoid or ellipsoidal organisms, often occurring in pairs, more plump and staining more deeply with the Giemsa stain than Rickettsia prowazekii. Da Rocha-Lima gives their size as 0.2 to 0.4 micron by 0.3 to 0.5 micron. In lice appear as short rods, frequently in pairs and often bipolarly stained. Non-mo- tile. Stain reddish-violet with the Giemsa stain. Gram-negative. Occur extra- FAMILY RICKETTSIACEAE 1095 cellularly in the region of the epithelial lining of the gut of the louse. Cultivation: Has not been cultivated in tissue culture or any cell-free medium, though Rickettsia pediculi, considered by some identical with Rickettsia quintana, has been cultivated on human and horse blood agar. Pathogenicity : Pathogenic for man, causing recurrent fever. No strain has been definitely established in laboratory animals. Immunology : Partial immunity is pro- duced after an attack of the disease. The disease is characterized by relapses which may occur as long as two years after the initial attack. Distinctive characteristics: The or- ganism resists a temperature of 60°C moist heat for 30 minutes or a dry heat at 80°C for 20 minutes. It resists desic- cation in sunlight for 4 months. It is filterable under certain conditions but not when in plasma or serum. It is present in filtrates of infected vaccine sediments and excrements of infected lice. Source : Seen in lice fed on trench fever patients by Topfer (Miinch. med. Wchnschr., 61, 1916, 1495). Habitat: The epithelial lining of the gut of the body louse (Pediculus humanus var. corporis) where they occur extra- cellularly, and Pedicuhis capitis. The virus is not transmissible through the ova. May be the etiological agent of trench fever (Wolhynian fever, shin bone fever, five-day fever). Appendix II : Additional named species are included in Chapter V, Rickettsiae, in Steinhaus, Insect Microbiology. Ithaca, 1946, 304-328. Some differ morphologically and tinctorially from typical rickettsiae, some are not asso- ciated with an arthropod vector, some have been incompletely studied and described, some have been cultivated in cell-free media. Pending the completion of further studies involving possible cultivation in fertile eggs, the determina- tion of biological properties, and ade- quate comparative immunological and serological studies, no attempt is made to classify these organisms. The de- scriptions are condensed from those given by Steinhaus : Ehrlichia (Rickettsia) kurlovi Mosh- kovsky. (Compt. rend. Soc. Biol., Paris, 126, 1937, 379; Ehrlichia kurlovi Mosh- kovsky, Uspekhi Souremennoi Biologii (Russian) (Advances in Modern Bi- ology), 19, 1945, 12.) Found in the monocytes of guinea pigs. Described by Kurloff in 1889 as inclusions in the mono- nuclear cells of guinea pigs and other animals. These became known as Kur- loff bodies. However, the parasitism of these bodies is questionable. Rickettsia avium Carpano. (Riv. Pat. Comp., Jan .-Feb., 1936, 1.) Minute bodies in the leucocytes and tissue cells of a bullfinch {Pyrrhula europea) brought to Egypt from Germany. Donatien and Lestoquard (Arch. Inst. Pasteur Algerie, 15, 1937, 142) suggested that this organism might have been that of psitta- cosis. Rickettsia bovis Donatien and Lesto- quard. (Donatien and Lestoquard, Bull. Soc. Path. Exot., 29, 1936, 1057; Ehrlichia bovis Moshkovsky, Uspekhi Souremen- noi Biologii (Russian) (Advances in Modern Biology), 19, 1945, 18.) Con- cerned in a disease of cattle which is transmitted by an unidentified tick of the genus Hyalomma. The organism occurs in circular or round-angled poly- gonal masses which consist of a large number of tightly pressed, minute spherical granulations. These masses are situated in the cytoplasm of various monocytes. The organism causes a rela- tively light febrile disease in cattle, and an inapparent infection in sheep and fever in monkeys. Rickettsia canis Donatien and Lesto- quard. (Donatien and Lestoquard, Bull. Soc. Path. Exot., 28, 1935, 418; Ehrlichia (Rickettsia) canis Moshkovsky, Compt. 1096 MANUAL OF DETERMINATIVE BACTERIOLOGY rend. Soc. Biol., Paris, 126, 1937, 382; Ehrlichia canis Moshkovsky, Uspekhi Souremennoi Biologii (Russian) (Ad- vances in Modern Biology), 19, 1945, 18.) Moshkovsky selects this species as the type species of the subgenus Ehrlichia Moshkovsky (loc. cit.). Found in dogs used for experimental purposes in Al- geria. Appears to be transmitted natu- rally by the dog tick (Rhipicephalus sanguineus). All active stages of the tick transmit the organism and it passes iutraovarially from the female to the larvae of the next generation. The or- ganisms are generally spherical in shape and can be seen in the circulating mono- cytes. The infection causes a serious and often fatal illness in dogs. Small laboratorj' animals are not susceptible to the disease. Rickettsia conjunctivae, Rickettsia con- junctivae bovis and Rickettsia conjunc- tivae gain, see Family III, Chlamydozo- aceae. Rickettsia ctenocephali Sikora. (Arch. Schiffs- u. Tropenhyg., 22, 1918, 442.) Found in cat fleas (presumably Cteno- cephalides felis) on the surface of the organs in the body cavity and in the coe- lomic fluid. Two forms were found which might be two species, one resembling Rickettsia pediculi and the other Rickett- sia nielophagi. Hertig and Wolbach (Jour. Med. Res., 44, 1924, 329) found Rickettsia ctenocephali to vary in size and shape from minute cocci to rather large, swollen, curved rods, staining reddish with the Giemsa stain. Rickettsia culicis Brumpt. (Ann. Para- sitol. Hum. et Comp., 16, 1938, 153.) Found in the stomach epithelium of mosquitoes {C ulex fatigans) 12 days after they had been fed on a patient carrying Microfilaria bancrofti. Thought to be pathogenic for the mosquito and possibly for man. Occurs in the form of small granules and more often as small bipolar rods. Stains with haemalum, erythro- sine-orange and toluidine blue. Gram- negative. Rickettsia dermacentrophila Steinhaus. (Pub. Health Repts., 57, 1942, 1375.) Found in all stages of the wood tick {Der- macentor andersoni). In the epithelial cells of the intestinal diverticula and other tissues of the tick, usually extra- cellularly but sometimes intracellularly. Xot seen in the nuclei of cells. Gram- negative and staining red with the Mach- iavello stain, and bluish-purple with the Giemsa stain. Stains less deeply with ordinary bacterial stains than most bacteria. Resembles Rickettsia rickett- sii morphologically but is slightly larger. Not pathogenic for laboratory animals or for some of the natural hosts of Derma- centor andersoni. Rickettsia hirundinis Cowdry. (Jour. Exp. Med., 37, 1923, 431.) An organism observed by Arkwright, Atkin and Bacot (Parasitology, 13, 1921, 27) in the tissues of Cimex hirundinis which is probably the same organism to which Cowdry re- ferred as Rickettsia hirundinis. Con- sidered by Steinhaus as a nomen nudum. Rickettsia kairo da Rocha-Lima. (Cairo rickettsia, Arkwright and Bacot, Brit. Jour. Exper. Path., 4, 1923, 70; da Rocha-Lima, in Kolle and Wasserman, Handb. d. path. Mikroorg., 3 Aufl., 8, 1930, 1361 . ) Resembles Rickettsia rocha- limae and Rickettsia prowazekii. Rickettsia lectularia Arkwright, Atkin and Bacot. (Parasitology, 13, 1921, 27.) Found in the gut of the bedbug {Cimex lectularius) as filamentous and rod- shaped organisms. It seems probable that all bedbugs harbor the organism and it is also present in the developing ova. The location is intracellular. Very pleomorphic, ranging from small coccoid forms to thread-like forms. The small coccoid and diplococcoid forms stain deep purple with the Giemsa stain, while bacillary, lanceolate and thread forms stain more red than purple with the Giemsa stain. Not infective for small laboratory animals or for man. Rickettsia linognathi Hindle. (Para- sitology, 13, 1921, 152.) Found in the FAMILY RICKETTSIACEAE 1097 alimentary tract of the goat louse (Lino- gnathus stenopsis). Resembles Rickett- sia trichodectae morphologically and oc- curs only extracellularly in the lumen of the gut. Rickettsia melophagi Noller. (Arch. Schiffs- u. Tropenhyg., 21, 1917, 53.) Found upon and in the cuticular layer covering the epithelium of the midin- testine of the sheep tick (Melophagus ovinus). Occurs characteristically in pairs of fairly uniform size, coccoid and sometimes rod-shaped. Gram-negative but stains fairly well with carbol-fuchsin and gentian violet. Stains deep purple with Giemsa's method and bright red with Machia Velio's method. Has been cultivated on non-living culture media, a glucose-blood-bouillon agar medium. The ability of Rickettsia melophagi to infect sheep has been the subject of contradictory claims. Small laboratory animals seem not to be susceptible. Rickettsia ovina Lestoquard and Dona- tien. (Lestoquard and Donatien, Bull. Soc. Path. Exot., 29, 1936, 108; Ehrlichia ovina Moshkovsky, Usphekhi Soure- mennoi Biologii (Russian) (Advances in Modern Biology), 19, 1945, 18.) Found in the blood of diseased sheep from Turkey and Algeria. The organisms oc- cur as minute coccoid granules, grouped in masses and present only in the mono- cytes and never in endothelial cells. They stain uniformly dark red with the Giemsa stain but did not stain with the Castaiieda technic. Infected ticks {Rhi- picephalus bursa) are thought to be the vectors. Rickettsia pisces Mohamed. (Ministry Agr., Egypt., Tech. Sci. Serv. Bull. 214, 1939, 6 pp.) In the monocytes and plasma of the blood of a fish {Tetraodon fahaka) showing necrotic ulcers on its head and both sides of the body. The heart, liver and intestines showed le- sions. The organisms were minute coc- coid forms varying from 0.2 to 0.4 micron in diameter and frequently occurring in pairs. Rickettsia rocha-liynae Weigh (Prz- glad. Epidemj., 1, 1921, 375.) Occurs in lice (Pediculus humanus) but is ap- parently non-pathogenic either to lice or to vertebrates. Larger and more pleo- morphic than Rickettsia prowazekii. In smears or sections of the gut of lice, Rickettsia rocha-limae occurs in agglomer- ated masses, grouped like staphylococci. They occur both extracellularly and in- tracellularly and stain more deeply than Rickettsia prowazekii. Weigl claims to have cultivated this species on artificial culture media under anaerobic condi- tions. Not pathogenic for laboratory animals or man. Rickettsia suis Donatien and Gayot. (Bull. Soc. Path. Exot., 35, 1942, 324.) Causes a disease in swine, the pathology of which resembles heartwater of rumi- nants. See Genus III, Coivdria, Family Rickettsiaceae. Rickettsia trichodectae Hindle. (Par- asitology, 13, 1921, 152.) In the species of biting lice {Trichodectas pilosus) which may be found on horses. This insect does not suck blood. The or- ganisms occur extracellularly in the ali- mentary tract of the louse. The aver- age size is 0.3 to 0.5 by 0.5 to 0.9 micron and occasionally longer forms occur. Rickettsia weigli Mosing. (Arch. Inst. Pasteur, Tunis, 25, 1936, 373.) Con- cerned in an epidemic disease which broke out in 1934 among employees of the Institute of Biology in Lwow who were engaged in feeding supposedly un- infected lice on their persons. Mosing and others have suggested the possibility that this rickettsia may be an extreme mutant of Rickettsia pediculi. Small coccoid to rod-shaped organisms staining well with the Giemsa stain, usually slightly longer than Rickettsia prowa- zekii. In the louse {Pediculus hu- manus), the rickettsiae occur extracellu- larly in the intestinal lumen forming a layer covering the surface of the epithe- lial lining. Not pathogenic for the louse as is Rickettsia prowazekii and Rickettsia 1098 MANUAL OF DETERMINATIVE BACTERIOLOGY rocha-liinae. It causes a febrile illness in man in which relapses occurred 3 to 5 times as in trench fever. Rickettsia weigli was agglutinated by convalescent sera but not by sera from typhus pa- tients. Convalescent sera gave no posi- tive Weil-Felix reaction. Wolbachia pipientis Hertig. (Rickett- sia of Culex pipiens, Hertig and Wolbach, Jour. Med. Res., U, 1924, 329; Hertig, Parasitology, 28, 1936, 453.) This is the type species of the genus Wolbachia Hertig {loc. cit ). Found in the ovaries or testes of the mosciuito, and present in all stages of the mosquito's development. The outstanding morphological charac- teristic of the organism is great pleomor- phism. Minute coccoids and short rods may be considered typical, but the usual microscopic field consists of various shapes and sizes. Som.e forms show bipolar staining with the Giemsa stain. The organism is a harmless parasite of the mosquito. Laboratory animals are apparently not susceptible. The following unnamed rickettsiae isolated from animals or seen in anihials are included in Steinhaus' list of rickett- siae (Insect Microbiology. Ithaca, 1946, 344): A rickettsia was isolated by Parker, Kohls, Cox and Davis (Pub. Health Rept., 5Jf, 1939, 1482) from a tick {Avi- hlyomma maculatiun) . It is pathogenic for guinea pigs and the disease is referred to as the maculatum- disease. There is complete cross immunity in guinea pigs between this infection and Rocky Mountain spotted fever and boutonneuse fever, but it differs from these diseases in some particulars. A rickettsia-like organism was isolated from the reduviid bug {Triatoma ruhro- fasciata) by Webb (Parasitology, 32, 1940, 355). It was pathogenic for some laboratory animals and was maintained in guinea pigs for 5 passages. The rickettsiae were transmissible to the next generation through the egg of the reduviid bug. A spotted fever type of rickettsia was isolated by Anigstein and Bader (Texas Repts. Biol. Med., 1, 1943, 105) from the dog tick {Rhipicephalus sanguineus) taken from normal dogs. It was patho- genic for rabbits and guinea pigs. A rickettsia was isolated by Anigstein and Bader (Texas Repts. Biol. Med., 1, 1943, 298, 389) from ticks {Amhlyomma americanum) collected in Texas. They believed it to be the cause of bullis fever. Rickettsiae were observed by Enigh (Berl. u. Mlinch. Tierarztl. Wchnschr., 1942, 25) in the leucocytes of a bison calf. No arthropod was associated with this rickettsia. A rickettsia-like agent pathogenic for guinea pigs was reported by Tatlock (Proc. Soc. Exp. Biol, and Med., 57, 1944, 95). The animals had been in- jected with blood from a patient with "pre tibial" fever. No arthropod vector was indicated. Three species of rickettsia-like or- ganisms isolated from the wood-tick (Dermacentor andersoni) are described by Noguchi (Jour. Exper. Med., 43, 1926, 518-521). These were named Ba- cillus rickettsiformis, Bacillus pseudo- xerosis and Bacillus equidistans. All could be cultivated on cell-free media and none was pathogenic for laboratory animals. Appendix III: Unnamed rickettsia- like organisms seen in the tissues of insects. Hertig and Wolbach (Jour. Med. Res., 44, 1924, 329) list sixteen species of arachnids and twenty-three species of insects which are hosts to rickettsiae or rickettsia-like organisms. Wolbach (Jour. Amer. Med. Assoc, 84, 1925, 723) reports hosts of non-patho- genic rickettsiae which include fourteen species of arachnids (ticks, mites and spiders) and twenty-two species of in- sects distributed in nine orders, including FAMILY RICKETTSIACEAE 1099 numerous non-blood-suckiug insects as well as lice and ticks. Cowdry (Arch. Path, and Lab. Med., 2, 1926, 59) lists seven species of arach- nids and twenty-four species of insects which are hosts to non-pathogenic rick- ettsiae. Buchner, P. (Tier und Pfianze in Sym- biose. Gebx-iider Borntraeger, Berlin, 1930, 900 pp.) Through the text, and particularly on pages 300-664, the rickettsia-like and bacterium-like micro- organisms occurring intracellularly in insects and other small animals are dis- cussed, principally from the viewpoint of the biologist. Paillot, A. (L'infection chez les In- sects, Paris, 1933, 535 pp.). Concerned principally with bacterial infections of insects, but also includes information in intracellular symbiotes and rickettsia- like and bacterium -like microorganisms. Steinhaus, Edward A. (Insect micro- biology. Ithaca, 1946, 188-255.) Intra- cellular bacterium-like and rickettsia- like symbiotes are discussed. 1100 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY II. BARTONELLACEAE GIESZCZYKIEWICZ.* (Bull. Intern. Acad. Polon. Sci., Classe Sci. Math. Nat., B (I), 1939, 9-30.)** Small, often pleomorphic, rod-shaped, coccoid, ring-shaped, filamentous and beaded micro-organisms, staining lightly with aniline dyes, but well with Giemsa's stain. Gram-negative. Parasites of the erythrocytes in man and other vertebrates. Known to be transmitted by arthropod vectors in some cases. The causative organisms of bartonellosis in man, haemobartonellosis, grahamellosis and eperythrozoonosis in the lower animals. Differ from the protozoa that also parasitize erythrocytes in that the entire parasite stains with no differentiation into cytoplasm and nucleus. Key to the genera of family Bartonellaceae. 1. Parasites of the erythrocytes and of fixed tissue in man. Genus I. Bartonella, p. 1100. 2. Parasites of the erythrocytes of lower mammals, increased in susceptible animals by splenectomy. Eradicated by arsenicals. Genus II. Haemobartonella, p. 1102. 3. Parasites of the erythrocytes of lower mammals. Not increased in susceptible animals by splenectomy. Not eradicated by arsenicals. Genus III. Grahamclla, p. 1109. 4. Blood parasites, found on the erythrocytes and in the plasma of lower mammals. Appear as rings, coccoids and short rods. Splenectomy activates latent infections. Genus IV. Eperythrozoon, p. 1111. Genus I. Bartonella Strong, Tyzzer and SeUards. {Bartonia Strong, Tyzzer, Brues, SeUards and Gastiaburii, Jour. Amer. Med. Assoc, 61, 1913, 1715; not Bartonia Muhlenberg, in Willdenow, Neue Schrift Ges. Nat. Fr., Berlin, 3, 1801, 444; not Bartonia Sims, Bot. Mag., 1804; not Bartonia Grossman, Essais de Paleoconchologie Comparee, 4me Livr., Paris, 1901; Strong, Tyzzer and SeUards, Jour. Amer. Med. Assoc, 64, 1915, 808; emend. Tyzzer and Weinman, Amer. Jour. Hyg., 30 B, 1939, 143.) Named for A. L. Barton who des- cribed these organisms in 1909. Parasites of the erythrocytes which also multiply in fixed tissue cells. On the red blood cells in stained films, they appear as rounded or oval forms or as slender, straight, curved or bent rods occurring either singly or in groups. Characteristically in chains of several segmenting organisms, sometimes swollen at one or both ends and * Prepared by Dr. Ida A. Bengtson (retired), National Institute of Health, Bethesda, Maryland and Dr. David Weinman, Parasitologist to the 1937 Harvard Expedition to Peru, Boston, Mass., April, 1947. ** Partial syn. Anaplasmidae has been proposed as a family name to unite the four genera Anaplasma, Grahamella, Bartonella and Eperythrozoon by Neitz, Alexander and du Toit (Onderst. Jour. Vet. Sci. and An. Ind., 3, 1934, 268). Since the name is derived from Anaplasma, the nature of which is not fully understood and since these authors consider the 4 genera as belonging to the protozoan order Haemosporidia, it seems advisable not to consider this nomenclature for the present. The genus Anaplasma (parasites of the red blood cells of cattle) created by Theiler (Transvaal Govt. Vet. Bact. Kept. 1908-9, 7-64, 1910) consists of two species Anaplasma mar- ginale and Anaplasma centrals. Recent workers are inclined to consider them to be bacterial in nature as they do not show a differentiation into cytoplasm and nucleus. FAMILY BA RTONELLACEAE 1101 frequently beaded (Strong et al., loc. cit., 1913), without a distinct differentiation of nucleus and cytoplasm. In the tissues they are situated within the cytoplasm of endothelial cells as isolated elements and grouped in rounded masses. These para- sites occur spontaneously in man and in arthropod vectors, are endowed with inde- pendent motility, reproduce by binary fission, and may be cultivated by unlimited serial transfers on cell-free media. One species has been recognized. It is known to be established only on the South American continent and perhaps in Central America. Human bartonellosis may be manifested clinically by one of the two syn- dromes constituting Carrion's disease (Oroya fever or verruga peruana) or as an asymptomatic infection (definition by Strong, Tyzzer and Sellards emend. Tyzzer and Weinman (in Weinman, Trans. Amer. Philosoph. Soc, N.S., 33, pt. 3, 1944, 246). The tj'pe species is Bartonella bacillijormis (Strong et al.) Strong et al. 1. Bartonella bacilliformis (Strong, Tyzzer, Brues, Sellards and Gastiaburu) Strong, Tyzzer and Sellards. {Bartonia bacilliformis Strong et al.. Jour. Amer. Med. Assoc, 61, 1913, 1715; Bartonella hacillijormis Strong, Tyzzer and Sellards, Jour. Amer. Med. Assoc, 64, 1915, 808; eynend. Tyzzer and Weinman, Amer. Jour. Hyg. 30(B), 1939, 143; also see Weinman, Trans. Amer. Philosoph. Soc, X.S., 33, pt. 3, 1944, 246. Partial or com- plete synonyms: Bartonella cocoide (sic) Hercelles, Ann. de Fac de Med., Lima, 9, 1926, 231; Bartonella peruviana Esco- mel, Bull. Soc path. Exot.. 22, 1929, 354; Eperthyrozoon noguchii Lwoff and Vaucel, Compt. rend. Soc Biol., Paris, 103, 1930, 975.) From Latin bacillus, rod and forma, shape. Small, pleomorphic organisms, show- ing greatest morphological range in the blood of man, appearing as red-violet rods or coccoids situated on the red cells, when stained with Giemsa's stain. Ba- cilliform bodies are the most typical, measuring 0.25 to 0.5 by 1 to 3 microns. Often curved and maj' show polar en- largement and granules at one or both ends. Rounded organisms measure about 0.75 micron in diameter and a ring- like variety is sometimes abundant. On semi-solid media a mixture of rods and granules appear. The organisms may occur singly or in large and small, ir- regular dense collections, measuring up to 25 microns or more in length. Puncti- form, spindle-shaped and ellipsoial forms of the organism occur, varying in size from 0.2 to 0.5 by 0.3 to 3 microns. Gram-negative and non-acid-fast. Stain poorly or not at all with the usual aniline dye stains, but satisfactorily with Romanowsky and Giemsa stains. Motile in the blood and in cultures. One to four unipolar flagella. Cultivation : Growth in semi-solid agar with fresh rabbit serum and rabbit hemoglobin and in semi-solid agar with blood of man, horse or rabbit with or without the addition of fresh tissue and certain carbohydrates, in other culture media containing blood, serum or plasma, Hun toon's hormone agar at 20 per cent, semi-solid gelatin media, blood-glucose- cystine agar, chorio-allantoic fluid and yolk sac of chick embryo. Gelatin not liquefied. No acid or gas n glucose, sucrose, galactose, maltose, fructose, xylose, lac- tose, mannose, mannitol, dulcitol, ara- binose, raffinose, rhamnose, dextrin, inu- lin, salicin and amygdalin. No action on lead acetate. Aerobic, obligate. Optimum temperature 28°C. Immunology : Natural immunity to infection has not been demonstrated in susceptible species. Acquired immunity apparent both during and after the dis- ease. Bartonellae from different sources appear to provoke similar responses. Bartonellae from Oroya fever protect 1102 MANUAL OF DETERMINATIVE BACTERIOLOGY against infection with organisms ob- mals, except rarely in an atypical form tained from verruga cases. in monkeys. Experimental verruga per- Serology : Immune sera fix complement uana has been produced in man, in a and agglutination of suspensions of number of species of monkeys and occa- Bartonella by sera from recovered cases sionally in dogs, has been reported. Source : Blood and endothelial cells Pathogenicity : Three forms of the dis- of lymph glands, spleen and liver of ease occur in man; the anemic (Oroya human cases of Oroya fever, fever), the eruptive (verruga peruana) Habitat: Blood and endothelial cells and mixed types of both of the other of infected man, probably also in sand forms. Experimental Oroya fever has flies (Phleboiomus verrucarum and Phle- not been successfully produced in ani- botomus noguchii). Genus II. Haemobartonella Tyzzer and Weinman. (Amer. Jour. Hyg., 30(B), 1939, 141.) From Greek haemos, blood and the generic name Bartonella. Includes parasites of the red blood cells in which there is no demonstrable multi- plication in the tissues and which do not produce cutaneous eruptions. They are typically rod- or coccoid-shaped, showing no differentiation into nucleus and cyto- plasm, occurring naturally as parasites of vertebrates, and are transmitted by arthro- pods. They are distributed over the surface of the erythrocj'tes, and possibly some- times within them. They stain well with Romanowsky type stains and poorly with other aniline dyes. Gram-negative. Not cultivated indefinitely in cell-free material. Rarely produce disease in animals without splenectomy, are markedly influenced by arseno therapy, and are almost all of world-wide distribution. The experimental host range is restricted, infectivity of a rodent species for other rodents being common, but for primates unknown. The type species is Haemobartonella muris (Mayer) Tyzzer and Weinman. Key to the species of genus Haemobartonella. I. The etiological agent of haemobartonellosis of the white rat. 1. Haemobartonella muris. II. The etiological agent of haemobartonellosis of the dog. 2. Haemobartonella canis. III. The etiological agent of haemobartonellosis of the vole. 3. Haemobartonella microtii. IV. The etiological agent of haemobartonellosis of the guinea pig. 4. Haemobartonella tyzzeri. V. The etiological agent of haemobartonellosis of cattle. 5. Haemobartonella bovis. VI. The etiological agent of haemobartonellosis of the buffalo. 6. Haemobartonella sturmanii. VII. The etiological agent of haemobartonellosis of the deer mouse. 7. Haemobartonella peromyscii. VIII. The etiological agent of haemobartonellosis of the gray-backed deer mouse. 7a. Haemobartonella peromyscii var. maniculati. IX. The etiological agent of haemobartonellosis of the short-tailed shrew. 8. Haemobartonella blarinae. X. The etiological agent of haemobartonellosis of the gray squirrel. 9. Haemobartonella sciurii. FAMILY BARTONELLACEAE 1103 1. Haemobartonella muris (Mayer) Tyzzer and Weinman. {Bartonella muris Mayer, Arch, f . Schiffs.- u. Tropen- hyg., 35, 1921, 151; Bartonella muris ratti Regendanz and Kikuth, Compt. rend. Soc. Biol., Paris, 98, 1928, 1578; Tyzzer and Weinman, Amer. Jour. Hj'g., 50(B), 1939, 143.) From Latin mus, muris, mouse. Slender rods with rounded ends, fre- quently showing granules or swellings at one or both extremities, and dumbbell, coccoid or diplococcoid forms. May oc- cur individually, in pairs, or in short chains of 3 or 4 elements, and, when abundant, in parallel grouping. The rods measure 0.1 by 0.7 to 1.3 microns and as much as half the length of a red cell. The coccoids have a diameter of 0.1 to 0.2 micron. They have been found on and in the erythrocytes and in the plasma. Pre- ferred stains are those of the Roman- owsky type. With Giemsa's stain various investigators report an intense red coloration, a bluish tinge with dis- tinct pink shading, blue with purple granules. With Wright's stain, the or- ganisms stain bluish, with reddish gran- ules at the ends. With Schilling's me- thylene blue-eosin stain the organisms stain a bright red color with the erythro- cyte staining blue. They stain faintly with Manson's stain, pyronin-methyl green and fuchsin. Gram-negative. There is lack of agreement concerning visibility in the fresh state and motility. Various authors report Brownian move- ment, slow and sinuous motion in the red cell or rapid motion. Cultivation: Cultivated with difficulty and divergent results have been reported. Growth on various media reported (blood agar, agar with 2 per cent defibrinated rat blood, horse blood agar, N. N. X., Blutrosplatte of Wethmar, hormone agar with blood of rabbit, horse or man, asci- tic fluid agar, chocolate agar, semi-solid rabbit serum agar, semi-solid rabbit blood agar, Noguchi-Wenyon medium, defibrinated rat blood, glucose broth. Tarozzi broth, peptone water) but usu- ally growth was scant or could not be continued by transfer to the same me- dium or the organism isolated was non- infectious or the possibility of latent infections in the animal was not excluded. Best results are apparently obtained with semi-solid rabbit serum agar and semi- solid rabbit blood agar. No conclusive results have been re- ported in tissue culture. The organism has been cultivated on the chorio-allan- toic membrane of the chick embryo. Filterability : Non-filterable with Seitz or Berkefeld N filters. Immunology : No authentic case of true natural immunity in rats has been established. Acquired immunity occurs in (1) the latently-infected rat, (2) the infected rat after splenectomy and re- covery from the disease, the period of resistance corresponding to the duration of latency, (3) the non-splenectomized non-carrier rat following infection, (4) animals other than the rat following in- fection. Serology : No precipitins, thrombocj'to- barin, isoagglutinins, or cold hemolysins have been reported in the serum of ane- mic rats. Complement deviation and agglutination have been reported with sera from rabbits, rats and guinea pigs injected Avith cultures. Rabbits im- munized with cultures have given posi- tive Weil-Felix reactions with Proteus 0X19 and OXK and rat sera recovered from haemobartonellosis have given a positive Weil-Felix reaction and positive agglutination in low dilution with Rickett- sia prowazekii. Pathogenicity : Infected blood, liver suspension, defibrinated laked blood, washed red cells, plasma and hemoglo- binuric urine may produce infection by the subcutaneous, intravenous, intra- peritoneal or intracardiac routes. Slight, transient or no haemobartonellosis oc- curs in adult non-splenectomized haemo- bartonella-free albino rats, adult non-splenectomized albino rats of carrier stock, adult splenectomized rats pre- 1104 MANUAL OF DETERMINATIVE BACTERIOLOGY viously infected, until 15 weeks to 8 months after infection. Typical haemo- bartonellosis occurs in adult splenec- tomized haeniobartonella-free albino rats and in young non-splenectomized haemo- bartonella-free albino rats weighing 20 to 30 grams at 3 weeks. Variable results have been obtained by different investi- gators with wild mice, guinea pigs, rabbits, hamsters, pigeons and monkeys {Macacus rhesus and Macacus sp.). It is known to be infectious for wild rats, albino mice, rabbits and for two Palestin- ian rodents {Sphallax {Spalax correct designation) typhlops and Meriones tris- trami). Negative results have been re- ported in dogs, kittens, cats, sheep and various birds. Causes a definite and characteristic anemia without cutaneous eruption. Arsenical therapy : True sterilization of latent or recognized infection with organic arsenical compounds. Source: Blood of infected albino rats. Habitat : Ectoparasites such as the rat louse {Polyplax (Haematopinus) spinulosus), the flea (Xenopsylla cheo- pis) and possibly the bedbug {Cimex lectularius) . Also found in the erythro- cytes of susceptible animals. World wide in distribution. 2. Haemobartonella canis (Kikuth) Tyzzer and Weinman. {Bartonella canis Kikuth, Klin. Wchnschr., 1928, 1729; Tyzzer and Weinman, Amer. Jour. Hyg., SO(B), 1939, 151.) From Latin canis, dog. One of the most pleomorphic of the haemobartonellae, occurring as thin rods, straight or slightly curved, dumbbell- shaped organisms, dots, coccoids, or rings. Chains of rods, coccoids or rings occur. These consist of only one type of these forms or a mixture of types. The chains may be straight, curved, branched or annular. Variable in size. Round forms vary from 0.2 or 0.5 micron to the limit of visibility. Single rods are 0.2 by 1 to 5 microns, while the composite forms vary from 1 to 4 microns. Situation is epi-erythrocytic. Giemsa's fluid stains the organism red-violet, usually intensely. Methyl- ene blue used as a vital stain colors the organism distinctly. Gram-negative and non-acid-fast. Considered non-motile by most inves- tigators. Cultivation : Cultivation has not been demonstrated in semi-solid rabbit serum- agar mediuni nor in media containing serum of splenectomized dogs, N.N.N., Noguchi's medium for leptospira, blood broth, Chatton's medium covered with vaseline for Trichomastix. Filterability : Results equivocal. Immunology : The outstanding phe- nomena resemble those found in the rat infected with Haemobartonella muris. Pathogenicity : Splenectomy is essen- tial to infection accompanied by anemia in the dog. Negative results in splenec- tomized haemobartonella-free guinea pig, rat, rabbit, and monkey {Cercopithecus sabaeus). No infection or anemia in unoperated mice, white rats, young rab- bits, young dogs and young guinea pigs. The splenectomized cat has been found to carry the infection by serial passage. Arsenical therapy : Complete sterili- zation obtained by neoarsphenamine. Source: Erythrocytes of infected sple- nectomized dogs. Habitat: Found in dog fleas (Cteno- cephalus) and erythocytes of infected animals. Distribution wide-spread, the infection occurring spontaneoush'' in Europe, India, North and South Africa, North and South America. 3. Haemobartonella microtii Tyzzer and Weinman. (Tyzzer and Weinman, Amer. Jour. Hyg. ,? 30(B), 1939, 143; also see Weinman, Trans. Amer. Philosoph. Soc, N. S., 33, 1944, 312; questionable synonym Bartonella arvicolae Yakimoff, Arch. Inst. Past, de Tunis, 17, 1928, 350; Haemobartonella arvicolae Weinman, loc. FAMILY BARTOXELLACEAE 1105 cit., 290.) From the genus of voles, Microtus . In infected animal, morphology re- sembles that of Haemobartonella canis, the organisms occurring as rods, coccoids, filaments, club forms, ring forms and granular niasses. In addition to these forms there occur in Giemsa-stained blood films ovoids, diamond- or flame- shaped small forms as well as coarse seg- mented or unsegmented filaments up to 5 microns in length. Filaments may con- tain one or more rings, or may be com- posed in part or entirely of diamond- shaped, coccoid or ovoid elements, some- times in parallel rows. Rods often show intense bipolar staining. Coccoid forms, usually scattered, may occur as aggre- gates or clumps on the red cell, appar- ently embedded in a faint blue matrix. A pale blue veil-like substance may cover nearly half of one surface of the red cells and show at its border typical red-violet stained rods or filaments in the Giemsa-stained specimens. A bow- shaped arrangement of elements is char- acteristic. Organisms lie on the surface of the red cells. In cultures organisms are more uniform in morphology resembl- ing Bartonella bacilliformis. Individual organisms are fine rods, 0.3 by 1.0 to 2 microns, sometimes occurring in chains and often in clumps. Small round forms occur, measuring 0.5 micron in diameter, and occasionallj' round disk-like struc- tures. Cultivation: Growth in Noguchi's semi-solid serum agar 2 weeks after in- oculation with citrated or heparinized blood and incubated at 23°C shows as white rounded masses, measuring up to about 1 mm in the upper 15 mm of the tube. In tissue culture the organism grows in snaall, rounded compact masses within the cytoplasm of infected cells. Indefinite maintenance of the strains isolated on artificial media has not been possible. Pathogenicity : Splenectomized white mice and splenectomized laboratory reared voles are readily susceptible to infection. Xo marked anemia or any mortality in heavily infected animals. Splenectomized dogs, white rats and deer mice are not susceptible. Source and habitat : Erythrocytes of the vole {Microlus pennsijlvanicus pemi- sylvanicus) following splenectomy. The natural mode of transmission has not been determined though ticks or mites are suspected. 4. Haemobartonella tyzzeri (Wein- man and Pinkerton) Weinman: (Bar- tonella tyzzeri Weinman and Pinkerton, Ann. Trop. Med., 33, 1938, 217; Wein- man, Trans. Amer. Philosoph. Soc, 33, 1944, 314.) Named for Prof. Tyzzer who studied haemobartonellae. Single or composite rods from about 0.25 micron by 1.4 to 4.0 microns. Occa- sional granular swellings and enlarged poles. Short rods also occur averaging 0.2 to 0.3 by 0.8 micron and also round forms with diameters of 0.2 to 0.3 micron. Distributed irregularly in the red cells. Stain intensely red-violet with Giemsa's or May-Griinwald-Giemsa's so- lutions. Gram-negative. Cultivation : Initial cultures on Nogu- chi's semi-solid serum agar obtained ir- regularly. When incubated at 28°C, colonies appear as isolated white spheres about 1 mm in diameter in the upper 8 mm border of the medium. The clumps are composed of rods and granules, with larger round structures or disks occurring occasionally. Also cultivated on the Zinsser, Wei and Fitzpatrick modifica- tion of the ;Maitland medium. Pro- longed maintenance on semi-solid media has not been obtained. Pathogenicity: Splenectomized haemo- bartonella-free guinea pigs may be infected by blood or cultures injected subcutaneously or intraperitoneally. Splenectomized Haemobartonella muris- free rats are insusceptible when inocu- lated with infected guinea-pig blood. Macacus rhesus monkeys are also in- 1106 MANUAL OF DETERMINATIVE BACTERIOLOGY susceptible to inoculations of infected blood, tissue and cultures. Infection of the guinea pig is subclinical in its mani- festations, probably due to the small number of parasites in the blood. No definite anemia accompanies infection. Source and habitat : Erythrocj^tes of the Peruvian guinea pig iCavia por- cellus). Has also been encountered in Colombia but not in other parts of the world. Observed in latently infected animals only after splenectomy. The natural mode of transmission is unknown, though the flea may be a possible vector. 5. Haemobartonella bovis (Donatien and Lestoquard) Weinman. (Donatien and Lestoquard, Bull. Soc. Path. Exot., 27, 1934, 652; Bartonella sergenti Adler and Ellenbogen., Jour. Comp. Path, and Therap., 47, 193-i, 221; (?) Bartonella bovis Rodriguez, Rev. del Inst. Llorente, 13, 1935, 5; abst. in Bull. Inst. Past., 34, 1936, 1033; Weinman, Trans. Amer. Philosoph. Soc, N. S., 33, 1944, 308; Haemobartonella sergenti Weinman, loc. oil., 290.) From Latin bos, bovis, ox. Resembles Haemobartonella ynuris and //. canis. Occurs as rods, coccobacilli and cocci, singly, in pairs or short chains or groups of 10 or more elements. The rods measure 1.2 to 2 microns in length and are very slender. The coccobacilli occur singly or in pairs measuring 0.3 by 0.6 to 0.8 micron and the diameters of the cocci are about 0.3 micron. The parasite may occupy a central or marginal position on the red cell ; the number on a cell varying from 1 to 20. Not more than 20 per cent of the cells are parasi- tized. Using the Romanowsky stain, the or- ganisms stain similarly to the chromatin of Piroplasma spp. Source and habitat : In the blood of bulls in Algeria and in a non-splenectom- ized calf in Palestine. 6. Haemobartonella sturmanii Grin- berg. (Grinberg, Ann. Trop. Med., 33, 1939, 33; Weinman, Trans. Amer. Philo- soph. Soc, N. S., 33, 1944, 313.) Similar to Haemobartonella bovis and H. ca7iis in morphologj^ and staining properties. Occurs as rods, cocco-bacil- lary and coccoid forms, varying in length from 0.5 to 1.5 microns. The number of parasites per infected cell varies from 1 to 15 and they occur individually, scat- tered irregularly in clumps or sometimes in chains stretching across the cell. At the height of the infection more than 90 per cent of the cells are infected. Pathogenicity: Causes a temperature rise in buffaloes and slight anemia after direct blood inoculation. Splenectom- ized rabbits, hamsters and splenectom- ized calves inoculated with blood from infected buffaloes remained free of the parasite . Source and habitat: In the blood of buffaloes in Palestine. 7. Haemobartonella peromysciiTyzzer. (Proc Amer. Philos. Soc, 85, 1942, 377.) Named for the genus of deer mice, Pero- myscus. Occurs as delicate filamentous forms (which may be branched) on the red blood cells. These filaments may be- come beaded and give rise to a number of coccoids and rods from which ring forms may develop. Stains by Giemsa's method, but stain- ing process must be intense in order to demonstrate the organism. Pathogenicity : Infection transmissible to splenectomized white rats, white mice and voles, producing a more or less severe illness with anemia. Habitat: In the blood of the deer mouse (Peromyscus lencopns novabor- acensis) . 7a. Haemobartonella peromyscii var. maniculati Tyzzer. (Proc Amer. Philos. Soc, 85, 1942, 381.) Named for the species of mouse from which it was iso- lated . Occurs as rods and filamentous FAMILY BARTOXELLACEAE iio"; branched forms. Coarser filaments ap- pear to rise from rounded granules. Del- icate rods are preponderant, and minute coccoids appear occasionally. When transferred to the common deer mouse, coarser forms appear, including filaments and large coccoids, sometimes in chains. Pathogenicity : Pathogenic for gray- backed deer mice and the common deer mouse, but non-infective for splenectom- ized white mice. Habitat : Blood of the gray -backed deer mouse {Peromyscus maniculatus gra- cilis). 8. Haemobartonella blarinae Tyzzer. (Proc. Amer. Philos. Soc, 85, 1942, 382.) Named for the genus of shrews, Blarina. Extremeh^ pleomorphic with delicate rods and coccus-like forms, often occur- ring in chains which also contain larger elements which have a deeply stained, bead-like granule. In the early stages of infection they may occur as thick bands or filaments stretching over the red cells usually with a bead or granule. The bands take a bluish tint with Geimsa's stain, while the more delicate form stains a slat}' violet. The head is distinctly reddish. In the fully de- veloped infection, I'ods and filaments pre- dominate over rounded forms. The or- ganisms may be scattered on the svu'facc of the red cells or may form a dense cap which is intensel}^ stained. Rudimen- tary mycelia may be found radiating from a central portion and reddish stained material with ill-defined contours may occur at the ends of the mycelial branches. Pathogenicity: Pathogenic for the short-tailed shrew but not for deer mice or white mice. Causes anemia in the shrew. Habitat: In the blood of the short- tailed shrew {Blarina brevicauda) . 9. Haemobartonella sciurii Tyzzer. (Proc. Amer. Philos. Soc, 55, 1942, 385.) Named for the genus of gray squirrels, Sciwus. Very pleomorphic. Occurs as minute rods and filaments which are continuous or segmented. The rods and filaments vary in thickness, some are very uneven and some very coarse. Beaded chains ma3' develop from the thickened forms. The bead-like elements stain a dull reddish at the periphery with Giemsa's stain while the remainder is very faintly stained in contrast to the intensely stain- ing basophilic rods and filaments. Some of the rounded forms have the appearance of large, thick rings. Beads and rings may arise from slender deeply staining rods, simulating verj' closely spores within bacilli, though no germination of filaments from them has been observed. Pathogenicity : Slightl}' pathogenic for the gray squirrel, non-pathogenic for normal white mice. Habitat : Blood of the gray squirrel (Sciurus carolincnsis leiicotis). Appendix: Here are included (1) Hae- uiohartonella of undetermined specific rank, (2) Haentobartonella-like struc- tures in non-splenectomized mammals and in cold-blooded animals, (3) Invalid species (see Weinman, Trans. Amer. Philosoph. Soc, N. S., 33, 1944, 315). 1. Hameobartonellae of undetermined specific rank. Microorganisms are grouped according to host of origin and are considered to be haemobartonellae from the description of the original author; but the information furnished is not sufficient for further classification. Haemobartonellae similar to Haemo- harlonella muris in wild rats : Miis decu- manus, Mus norvegicus, Rattus rattiLs frugivorus, Mus rattus griseiventer, Mus rattus rattus, Mus sylvaticiis. In vari- ous rats; technical names not given. Haemobartonellae similar to Haemo- bartonella tniiris in albino mice. Schil- ling (Klin. Wchnschr., 1929, 55) separated the haemobartonella of the mouse from 1108 MANUAL OF DETERMINATIVE BACTERIOLOGY that of the rat and named it Bartonella muris musculi var. alhinoi {Haemobar- tonella muris musculi var. albinoi Wein- nmn, loc. cit., 290). Haemobartonellae similar to Haemo- barlonella muris in other mammals : Haemobarionella glis glis (Kikuth) Wein- man {Bartonella glis glis Kikuth, Cent, f. Bakt., I Abt., 123, 1931, 356; Weinman, loc. cit., 317) in dormice (Glis glis). Haemobartonella opossum (Regendanz and Kikuth) Weinman {Bartonella opos- sum Regendanz and Kikuth, Arch. f. Schiffs- u. Tropenhyg., 32, 1928, 587; Weinman, loc. cit., 290) in the marsupial rat {Metachirus opossum) and in the opossum {Didelphys didelphys) . Haemobartonella spp. in Lophuromys ansorgei, in Lophuromys laticeps, in Oenomys bacchante editus, in Praeomys jacksoni, in Arvicanthus striatus, in deer mouse {Perojnyscus leucopus novabora- censis), in Chinese hamsters {Cricetulus griseus, Cricetulus griseus fumatus), in Apodemus agrarius a.ud Phodopus praedi- lectus), and in squirrels {Sciurus vulgaris). Mixed infections, including haemobar- tonellae are found in jerboa, the gerbille and various rodents (see Weinman, loc. cit., 317-319). 2. Haemobartonella-like structures in non-splenectomized mammals and cold- blooded animals. Various bodies whose proper classifica- tion in the genus Haemobartonella has not been established (Weinman, loc. cit., 319) In non-splenectomized mammals : Bartonella melloi Yakimoff and Raste- gaieff. Bull. Soc. Path. Exot., 2^, 1931, 471 {Haemobartonella melloi Weinman, loc. cit., 290) in the ant eater {Manis pentadactijla) . Bartonella pseudocebi Pessoa and Prado, Rev. biol. e hyg., 1, 1927, 116 {Haemobartonella pseudocebi Weinman, loc. cit., 290) in the monkey {Pseudoce- bus apella). Bartonella rocha-limai Fariaand Pinto, Compt. rend. Soc. Biol., Paris, 95, 1926, 1500 {Haemobartonella rocha-limai Wein- man, loc. cit., 290) in the bat {Hemi- derma brevicauda). Bartonella sp. in the rat {Rattus rufes- cens) and Bartonella sp. in the dormouse {Myoxus glis). In cold-blooded animals: Bartonella pavlovskii Epstein, All Union. Inst. Exper. Med., Moscow, 1935, 398, see Ray and Idnani, Indian Jour. Vet. Sci. and Animal Husb. 10, 1940, 259, {Haemobartonella pavlovskii, Weinman loc. cit., 290) in the lamprey {Petromyzon marinus) . Bartonella nicollei Yakimoff, Arch. Inst. Pasteur Tunis, 17, 1928, 350 {Hae- mobartonella nicollei Weinman, loc. cit., 290) in the brochet {Esox lucius). Bartonella ranarum da Cunha and Munez, Compt. rend. Soc. Biol., Paris, 97, 1927, 1091 {Haemobartonella ranarum Weinman, loc. cit., 290) in the frog {Lep- todactylus ocellatus). This is pi'obably identical with Bartonella batrachorum Zavattari, Boll. d. Soc. ital. biol. sper., 6, 1931, 121 {Haemobartonella batracho- rum Weinman, loc. cit., 290) from the same species. Bartonella sp. in the gecko {Platydac- tylus mauritanicus) , Bartonella sp. in the lizard {Lacertilia sp.), Bartonella sp. in the lizard {Tropidurus peruvi- anus), Bartonella sp. in the tench {Tinea tinea) and Bartonella sp. in the tortoise {Testudo graeca). 3. Invalid species: Bartonella caviae Campanacci, Ateneo parmense, 1, 1929, 99 {Haemobartonella caviae Weinman, loc. cit., 290) from the guinea pig. Bartonella ukrainica Rybinsky, Rev. Microbiol, epidem. et parasitol., 8, 1929, 296 {Haemobartonella ukrainica Weinman, loc. cit., 290) from the guinea pig- Weinman {loc. cit., 314) states that the parasitism of these structures was not proven and no illustrations are fur- nished by the authors. FAMILY BARTONELLACEAE 1109 Genus III. Grahamella Brumpt. (Brumpt, Bull. Soc. Path. Exot., J^, 1911, 514; Grahamia Tartakowsky, Trav. IX-^ Cong. Int. Med. Vet., 4, 1910, 242; not Grahamia Theobald, Colonial Office, Misc. Pub. No. 237, 1909.) Named for G. S. Graham-Smith who discovered the parasite in the blood of voles. Parasites occurring within the erythrocytes of the lower mammals which morpho- logically bear a resemblance to Bartoyiella, but which are less pleomorphic, more plump, and more suggestive of the true bacteria. They stain more deeply than bar- to nellae with Giemsa's stain, stain lightly with aniline dyes and with methylene blue. They are Gram-negative, non-acid-fast and non-motile. Splenectomy has no effect on the source of infection e.xcept in rats. They are non-pathogenic and not affected by arsenicals. Several species have been cultivated on cell-free media.* The etio- logical agent of grahamellosis of rodents and some other vertebrates. The type species is Grahamella talpae Brumpt. 1. Grahamella talpae Brumpt. (Bull. Soc. Path. Exot., 4, 1911, 514.) Named for the genus of moles, Talpa. Long or short rods of irregular contour lying within the red blood cells, many with a marked curve, often near one of the extremities. One or both ends of the longer forms enlarged, giving a wedge- or club-shaped appearance. Some of the medium-sized forms definitely dumbbell - shaped, small forms nearly round. With Giemsa's stain, the protoplasm of the organism stains light blue, with darker areas at the enlarged ends. Dark staining areas of longer forms give the organism a banded appearance. Length varies from 0.1 to 1 micron. Parasites occasionally free in the plasma, but usually in groups. Most of the infected corpuscles contain between 6 and 20 parasites (Graham-Smith, Jour. Hyg., 5, 1905, 453). Pathogenicity: Pathogenic for moles. Appendix: In addition to Grahamella talpae Brumpt, descriptions of the following species occur in the literature. The list may not be complete and the validity of these species may be ques- tioned in some cases. Grahamella acodoni Carini. (Ann. Parasit., 2, 1924, 253.) From Acodon serre7isis, Brazil. Grahamella alactagae Tartakowsky. (Katalogue der Exponaten der Land- wirthschaftlichen Ausstellung (Rus- sisch), St. Petersburg, 1913.) From Alactaga saliens and Alaclaga aconitus in Transcaucasia and steppes of Astrakhan (Alactaga misspelled Alactoga). Quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 303. Grahamella arvalis Tartakowsky. (Katalogue der Exponaten der Land- wirthschaftlichen Ausstellung (Russisch) St. Petersburg, 1913.) From Microtus arvalis in Transcaucasia. Quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 304. Grahamella balfourii Brumpt. (Gra- hamella sp. Balfour, Rept., Wellcome Tropical Research Laboratory, 2, 1906, 97; Grahamella balfouri Brumpt, Bull. Soc. Path. Exot., Paris, 4, 1911, 517.) From the desert rat {Jaciilus jaculus) in the Sudan. * Tyzzer (Proc. Amer. Philos. Soc, 85, 1942, 375) finds that grahamellae isolated in culture show a close relationship to Streptobacillus moniliformis {Actinomijces muris) and proposes the inclusion of the genus Grahamella in the family Actinomy- cetaceae. The latter relationship appears to be very doubtful. 1110 MANUAL OF DETERMINATIVE BACTERIOLOGY Grahamella hlarinae Tyzzer. (Proc. Amer. Philos. Soc, 85, 1942, 370.) From the short-tailed shrew (Blarina brevi- cauda) in Massachusetts. Grahamella bovis Marzinowsky . (Med . Obosrenie, 1917, No. 1-2.) From the ox {Bos taunts) in Russia. Quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 304. Grahamella brumptii Ribeyro and del Aquilla. (Ann. Fac. Med., Lima, 1, 1918, 14-20.) From Desmodus rufus in Peru. Grahamella cams lupus Kamalow. (Cent. f. Bakt., I Abt., Orig., 138, 1933, 197.) From the wolf, Tiflis. Grahamella couchi Neitz. (Onderst. Jour. Vet. Sci. and An. Ind., 10, 1938, 29.) From the multimammate mouse (Masto- mys coucha) in South Africa. Grahamella criceii domestici Parzwa- nidze. (Das Material zum Hiimoparasi- tismus der Tiere bei Uns. Tiflis, 1925.) From Criceius doynesticus in Transcau- casia. Quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 304. Grahamella cricetuli Patton and Hindle. (Proc. Roy. Soc, London, BilOO), 1926, 387.) From Cricelulus griseus in China. Grahamella dschunkowskii Tartakow- sky. {Grahamia sp., Dschunkowsky and Luhs, Trav. IX. Cong. Internat. Med. Vet., 1909, 4, 1910, 242; Grahamia dschun- kowski Tartakowsky, 1910.) From the bat {Vcspertilio noctula) in Transcau- casia. Ref. to Tartakowsky quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 304. Grahamella dudtschcnkoi Yakimoff. {Grahamella sp., Dudtschenko, Cent. f. Bakt., I Abt., Orig., 74, 1914, 241; (7m- hamella dudtschenkoi Yakimoff, Arch. f. Protistenk., 66, 1929, 304.) From the hamster {Cricehdus sp.) in Transbaikal. Grahamella ehrlichii Yakimoff. (Gra- hamella ehrlichi Yakimoff, Arch. f. Pro- tistenk., 66, 1929, 305.) From the perch {Perca fluviatilis) in Russia. Grahamella francai Brumpt. {Gra- hamella sp. Franca, Arch. Inst. Bacter. Camara, Pestana, 3, 1911, 277; Graha- mella francai Brumpt, Precis de Parasi- tologic, 2eme ^d., 1913, 102.) From the, jumping rat {Eliomijs quercinus) in Portugal. Grahamella gallinarum Carpano. (Ann Parasit. hum. et comp., 13, 1935, 238.) From leghorn chickens in Egypt. Grahamella gerbilli Sassuchin. (Gra- hamia gerbilli Sassuchin, Arch. f. Pro- tistenk., 74, 1931, 526.) From Gerbillus tamaricinus in southeast Russia. Grahamella hegneri Sassuchin. (Gra- haynia hegneri Sassuchin, Arch. f. Pro- tistenk., 75, 1931, 152.) From Citellus pygmaeus in Russia. Grahamella joyeuxii Brumpt. (Gra- hamella sp., Joyeu.x, Bull. Soc. Path. Exot., Paris, 6, 1913, 614; Grahamella joyeuxi Brumpt, Precis de Parasitologic, 2eme ed., 1913, 102.) From Gol unda fallax and Mus ratlus in French Guinea. Grahamella merionis Adler. (Trans. Roy. Soc. Trop. Med., 24, 1930, 78.) From Meriones trislrami in Palestine. Grahamella microti Lavier. (Bull. Soc. Path. Exot., Paris, 14, 1921, 573.) From Microtus arvalis in France. Grahamella microti pennsylvanici Tyz- zer. fProc. Amer. Philos. Soc, 85, 1942, 366.) From the common vole {Microtus pennsylvanicus pennsylvanicus) in Massa- chusetts. Grahamella muris Carini. (Bull. Soc. Path. Exot., Paris, 8, 1915, 104.) From the house rat [Mus dccumanus) in Brazil. Grahamella muris musculi iberica Parzwanidze. (Das Material zum Hamo- parasitismus der Tiere bei uns. Tiflis, 1925.) From Mus musculus in Trans- caucasia. Quoted from Yakimoff, Arch. f. Protistenk., 66, 1929, 304. Grahamella musculi Benoit-Bazille. (Bull. Soc. Path. Exot., Paris, 13, 1920, 408.) From Mus ynusculus var. albinos in France. Grahamella ninae kohl-yakomovi Yaki- moff. (Bull. Soc. Path. Exot., Paris, 10, FAMILY EARTOXELLACEAE 1111 1917, 99.) From the hamster (Cricetus GrahameUa pipistrelli Markow. (Gra- phoca) in Transcaucasia. harnia pipistrelli Marlcow, Russian Jour. GrahameUa peromysci Tyzzer. (Proc. Trop. Med., 1926, No. 5, 52.) From the Amer. Philos. Soc, 85, 1942, 363.) From bat (Pipistrellus nalhusu) in Russia. the deer mouse (P. leucopusnovaboracen- GrahameUa rhesi Leger. (Bull. Soc. sis) m Massachusetts. p^^j^ E^^^., Paris, 15, 1922, 680.) From GrahameUa peromysci var. maniculati ,, , ,^r t, s • . ,^^ . T^, ., o the monlcey [Macacus rhesus) in Annam. GrahameUa sanii Cerruti . (Arch. Ital . Tyzzer. (Proc. Amer. Pliilos. Soc, 85, 1942, 365.) From the grav-backed deer . mouse (Peromyscus maniculatus) in ^ci. Med. Col.. //, 1930, 522.) From Massachusetts. Testudo graeca in Sardima. GrahameUa phyllotidis Tyzzer. (Proc. GrahameUa falassochelys Cerruti. Amer. Philos. Soc, 85, 1942, 371.) From (Arch. Ital. Sci. Med. Col., 12, 1931, 321.) the Peruvian mouse (PhyUotis darunni From Tallasochelys carelta in Sardinia. linatus). (Misspelled for Thalassochelys.) Genus IV. Eperythrozoon Schilling.* (Schilling. Klin. Wchnschr., 1928, 1854; Gyromorpha Dinger, Nederl. tijdschr. geneesk., 72, 1928, 5903.) From Greek meaning animal on red blood cell. Microscopic blood parasites found in the plasma and on the erythrocytes. They stain well with Romanowsky type dyes, and then appear as rings, coccoids or short rods, 1 to 2 microns in greatest dimension, staining bluish or pinkish violet. They show no differentiation of nucleus and cytoplasm. The organisms are not known to retain the violet in Gram's method or to be acid-alcohol-fast. Splenectomy ac- tivates latent infection. Not cultivated in cell-free media, .\rthropod transmission has been established for one species (Weinman, Trans. Amer. Philosoph. Soc, N.S. 53, pt. 3, 1944. 321). The type species is Eperythrozoon coccoides Schilling. Key to the species of genus Eperythrozoon. I. Etiological agent of eperj^throzoonosis of white mice. 1. Eperythrozoon coccoides. II. Etiological agent of eperythrozoonosis of sheep. 2. Eperythrozoon ovis. III. Etiological agent of eperythrozoonosis of cattle. 3. Eperythrozoon wenyonii. IV. Etiological agent of eperythrozoonosis of gray-backed deer mice. 4. Eperythrozoon varians. V. Etiological agent of eperythrozoonosis of voles and dwarf mice. 5. Eperythrozoon dispar. * This genus has been considered as belonging to the Protozoa by Neitz, Alexander and Du Toit (Onderst. J. Vet. Sci., 3, 1934, 268) and to the bacteria by Mesnil (Bull. Soc. Path, exot., 22, 1929, 531 and by Tyzzer (in Weinman, Trans. Amer. Philosoph. Soc, N.S., 33, pt. 3, 1944, 244). The evidence at hand favors the inclusion of this group among those organisms which are not protozoan in nature but which are closely related to bacteria. 1112 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Eperythrozoon coccoides Schilling. (Schilling, Klin. Wchnschr., 1928, 1854; Gyromorpha musculi Dinger, Nederl. tiidschr. geneesk., 72, 1928, 5905.) From Greek, coccus-shaped. In stained blood films these organisms appear as rings, coccoids and rods, the majority as rings of regular outline with clear centers. They are in the plasma and on the red cells. Measure 0.5 to 1.4 microns in greatest dimension. Stain pale red or reddish-blue with the Giemsa or the May-Griinwald-Giemsa technics. Gram-negative. Suggested methods of multiplication by binary fission, budding, development of small coccoidal to annular forms. Cultivation: Negative results. Immunology : Immunological state in animals that of the premunition type. Latent infection in mice which is made manifest by splenectomy. Pathogenicity : Pathogenic for white mice, rabbits, white rats, wild mice, usually in young animals or in splenec- tomized adults. Source : Blood of splenectomized white mice. Habitat: Blood of infected animals, mouse louse {Polyplax serrata) and probably other arthropods. 2. Eperythrozoon ovis Neitz, Alexan- der and Du Toit. (Neitz et al.. Address, Biological Society, Pretoria, Mar. 15, 1934; from Neitz, Onderst. Jour. Vet. Sci. and An. Ind., 9, 1937, 9.) From Latin ovis, sheep. Delicate rings approximately 0.5 to 1.0 micron in diameter though occasion- ally larger. In addition there are tri- angles with rounded angles, ovoid, comma, rod, dumbbell and tennis racket forms. Found supra-cellularly on the erythrocytes but often free. Colored pale purple to pinkish-purple with Giemsa's stain. Suggested mode of mul- tiplication by budding. Cultivation: Negative results. Immunology : Immunological state in sheep appears to be that of the premuni- tion type. Pathogenicity : Sheep, antelopes and probably goats and splenectomized calves are susceptible. Dogs, rabbits and guinea pigs are refractory. The distinc- tive feature of Eperythrozoon ovis is its ability to provoke illness in normal animals without resorting to splenec- tomy. Source : Blood of infected South Afri- can sheep. Habitat: Blood of infected animals. No ectoparasites found on sheep naturally infected, but an arthropod is suspected. 3. Eperythrozoon wenyonii Adler and Ellenbogen. (Adler and Ellenbogen, Jour. Comp. Path, and Therap., ^7, 1934 (Sept. 3), 220; see Bartonella wenyoni in appendix.) Named for Dr. C. M. Wen- yon, a student of these organisms. Morphologically similar to Eperythro- zoon coccoides. Coccoid and often vesi- cular, staining pale red with Giemsa's stain and varying from 0.2 to 1.5 microns in diameter. Multiplication seems to be by budding and fission, and by filamen- tous growths from the ring forms, suggest- ing resemblance to Hyphomyceles. Up to 50 or 60 parasites are found on one cell. These are arranged in irregular chains or in tightly packed groups. Cultivation not reported. Immunology : The organism creates a state of premunition and latent infection is made manifest by splenectomy. Pathogenicity : Cattle are susceptible, but sheep are not infected either before or after splenectomy. Source: Blood of infected cattle. Habitat : Blood of infected cattle, arthropod transmission not proven. 4. Eperythrozoon varians Tyzzer. (Proc. Amer. Philos. Soc, 85, 1942, 387.) From Latin varians, varying. FAMILY BARTONELLACEAE 1113 Occur in rings, coccoids of varying size, some very minute, bacillary forms. Many of the bacilliform elements show an unstained lens-like swelling, indicat- ing the formation of a ring within the substance of the rod. At the height of the infection most of the organisms are found in the plasma. Whenever an or- ganism comes in contact with a red cell, it stains intensely. Pathogenesis : Pathogenic for the gray- backed deer mouse (causing anemia) and for the splenectomized common deer mouse. Not pathogenic for splenectom- ized white mice. Habitat : Blood of the gray-backed deer mouse (Perotnyscus maniculatus gra- cilis). 5. Eperythrozoon dispar Bruynoghe and Vassiliadis. (Ann. de Parasitol., 7, 1929, 353.) Resembles Eperythrozoon coccoides in staining, distribution on the erythro- cytes and also in appearance except that circular disks with solid staining centers may greatly outnumber the ring forms. Found on the red blood cells and in the plasma. Size range that of Eperythro- zoon coccoides, also some larger ring forms. Cultivation: Not successful. Immunology: Infection is followed by premunition and latent infection is made manifest by splenectomy. Splenectom- ized rabbits premunized against E. coc- coides do not react to inoculation with E. dispar; if the latter is injected first, they do not react to E. coccoides. Infectivity : Infective for the European vole {Arvicola[Microtus] arvalis), the American vole (Microtus pennsylvanicus pennsylvanicus) , the dwarf mouse {Mus minutus), the rabbit, and Mus aconiys. Not infective for albino rats or albino mice. Source: Blood of infected animals. Appendix: 1) Species incompletely studied, Eperythrozoon spp. and Epery- ihrozoon-like structures (Weinman, Trans. Amer. Philosoph. Soc, N. S. S3, pt. 3, 1944,320). Eperythrozoon noguchii Lwoff and Vau- cel. (Bull. Soc. path, exot., 26, 1933, 397.) Probably not a valid species. Eperythrozoon perekropovi Yakimoff. (Arch. f. Protistenk., 73, 1931, 271.) Classification in genus Eperythrozoon questionable. Bartonella wenyoni Nieschulz. (Ztschr. f. Infektionskr., 53, 1938, 178.) Probably identical with Eperythrozoon wenyoni. If valid, Haemohartonella wenyoni. Possible human infection (Schiiffner, Nederl. tijdschr. v. geneesk., 73, 1939, 3778). 2) Animals infected with parasites which are definitely eperythrozoon-like but of uncertain specificity or which are eperythrozoon-like in some features but which can not be definitely classified generically : Jerboa sp. Kikuth. (Cent. f. Bakt., I Abt., Orig., 123, 1931, 356.) Arvicola arvalis Zuelzer. (Zuelzer, Cent. f. Bakt., I Abt., Orig., 102, 1927, 449; Kikuth, Ergebn. Hyg. Bakt., Im- munitatsforsch. u. Exper. therap., 13, 1932, 559.) Rattus rattns Schwctz. (Ann. Soc. beige de med. trop., U, 1934, 277.) Sciurus vulgaris Nauck. (Arch. f. Schiffs- u. Trop. -Hyg., 31, 1927, 322.) Leptodactylus pentadactylus Carini. (Compt. rend. Soc. Biol., Paris, 103, 1930, 1312.) 1114 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY III. CHLAMYDOZOACEAE IMOSHKOVSKY . * (Uspekhi Souremennoi Biologii (Russian) (Advances in Modern Biology), 19, 1945, 12.) Small, pleomorphic, often coccoid microorganisms usually with characteristic de- velopment cycle. Stain with aniline dyes. Gram -negative. Behave as obligate intracytoplasmic parasites. Have not been cultivated in cell-free media. Criteria adequate for classification lacking for more recentl}' isolated members. The attribu- tion of Genus III, Colcsiuta, either to Rickettsiaceae or to Chlamydozoaceae is still in doubt . Key lo the genera of family Chlamydozoaceae. I. Cells coccoid and with life cycle. A. Non-oultivatable in chicken embryonic tissues. Genus I. Chlamydozoon, p. 1114. B. Cultivatable in chicken embryonic tissues. Genus II. Miyagawanella, p. 1115. II. Cells pleomorphic. Genus III. Colesiota, p. 1119. Genus I. Chlamydozoon Ilalberstaedter and von Prowazek. fArb. a. d. kaiserl. Gesundheitsamte, 26, 1907, 44.) From Greek chlamydos, cloak and zoon, animal. Coccoid spherical cells with developmental cycle. Gram-negative. Intracyto- plasmic Iiabitat. Non-cultivatable in chicken embryonic tissues. Susceptible to sulfonamide and penicillin action. The type species is Chlamydozoon trachomatis Foley and Parrot. 1. Chlamydozoon trachomatis Foley dyes; blue or reddish-blue with the and Parrot. (Rickettsia trachomae Bus- Giemsa stain and red or blue, depending acca. Arch. Ophthalm., 52, 1935, 567; on the metabolic state, with the Macchi- Foley and Parrot, Arch. Inst. Past. d'Al- avello stain. Matrix of plaques gives a g4ne, 15, 1937, SS9; Rickettsia trachoma- strong reaction for glycogen. Non- ius Foley and Parrot, ic/e/n.) Named for motile, the disease, trachoma. Cultivation : Has never been culti- Coccoid bodies: Small microorganisms yot-gj 200 to 350 millimicrons in diameter form ^ i • i <- tr , ,. T • • , , 1- Immunological aspects: Has one or the elementary bodies. Initial bodies ^. . ■^.u ^ t .,,. . . ,. ^ 1 nwre antigens in common with or closely up to 800 millimicrons in diameter and , ,, ^ • ir- r X ir> • 1 f J Ml resembling one or more present m iii ?. va- plaques up to 10 microns also tound. All , * ,^ , • , larger forms encapsulated with substance Sawanella spp. Produces, in low con- derived either from the agent or from centrations, antibodies which fix comple- the cytoplasm of the parasitized cells. me^t with antigen from Miyagawanella Elementary body is the basic unit. lymphogranulomatis. Paired forms or clusters occur. Gram- Pathogenicity: Pathogenic for man, negative. Stains poorly with aniline apes and monkeys where it affects only * Prepared by Dr. Geoffrey Hake, The Squibb Institute for Medical Research, New Brunswick, New Jersey, September, 1946. FAMILY CHLAMYDOZOACEAE 1115 the corneaand conjunctiva causing highly Immunological aspects: As for C. tra- destructive lesions. chomatis. Chemotherapy : Susceptible to sulfon- Pathogenicity : Pathogenic for man, amides and penicillin. _ baboons and monkeys. Causes an acute conjunctivitis and, in man, an inflamma- Source : Found in scrapings of cornea or conjunctiva in cases of trachoma. Habitat: The etiological agent of tra- *'°" ^^ *^^ ^°^^^^ gemto-urinary tract, choma in man. Chemotherapy: Susceptible to sulfon- amides and penicillin. 2. Chlamydozoon oculogenitale Mosh- Source: Found in conjunctival exu- kovsky. (Moshkovsky, Uspekhi Sour- dates, and in exudates from infected emennoi Biologii, 19, 1945, 12.) From ^^^^^hra or cervix. Also present in con- Latin oculus, eye and genitalis, genital. , • , , , ^ , ,, , , 1 i • • i- taminated pools of water. Morphology and staining reactions : As for Chlamydozoon trachomatis. Habitat : The etiological agent of swim- Cultivation: Has never been culti- ming pool conjunctivitis, neonatal con- vated. junctivitis or inclusion conjunctivitis. Genus II. Miyagawanella Brumpt. (Ann. de Parasit., 16, 1938, 153.) Named for Prof. Miyagawa, the Japanese bacte- riologist, who first (1935) grew the type species in the chick embryo. Coccoid to spherical cells with a developmental cycle. Gram-negative. Intra- cytoplasmic habitat. Cultivatable in chicken embryonic tissues. Some species are susceptible to sulfonamide or penicillin action. The type species is Miyagawanella lymphogranulotnatis Brumpt. Key to the species of genus Miyagawanella. I. The etiological agent of lymphogranuloma venereum, lymphogranuloma in- guinale, climatic bubo, and esthiomene in man. 1. Miyagawanella lymphogranulotnatis. II. The etiological agent of psittacosis or parrot fever. 2. Miyagawanella psittacii. III. The etiological agent of ornithosis (Meyer). 3. Miyagawanella ornithosis. IV. The etiological agent of one type of viral pneumonia. 4. Miyagawanella pneumoniae. V. The etiological agent of mouse pneumonitis (Gonnert). 5. Miyagawanella bronchopneumoniae . VI. The etiological agent of feline pneumonitis (Baker). 6. Miyagawanella felis. VII. The etiological agent of Louisiana pneumonia. 7. Miyagawanella louisianae. VIII. The etiological agent called the Illinois virus, the cause of one type of viral pneumonia. 8. Miyagawanella illinii. 1116 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Miyagawanella lymphogranuloma- tis Brumpt.(Biumpt, Ann. de Parasit., 16, 1938, 153; Ehrlichia lymphogranulo- matosis Maui'o, (Reference not found.) Named for the disease, lymphogranu- loma. Ooccoid bodies : Small microorganisms 200 to 350 millimicrons in diameter form the elementary bodies. Initial bodies up to 1 micron and plaques up to 10 microns also found. All larger forms encapsulated with a substance derived either from the agent or from the cyto- plasm of parasitized cells. Elementary body is the basic unit. Paired forms or clusters occur. Gram-negative. Stain with aniline dyes, purple with the Giemsa stain and red or blue, depending on metabolic state, with the Macchiavello stain. Matrix of the plaque does not give the reaction for glycogen. Non-motile. Filterability : Passes through Chamb- erland Lo and L3, Berkefeld V and N and sometimes through Seitz EK filters. Cultivation : In plasma tissue cultures of mammalian cells, in mammalian cells on agar, in the chorio-allantoic membrane or particularly in the yolk sac of the chicken embryo but has not been culti- vated in the allantoic sac. Optimum temperature 37°C in tissue cultures, 35°C in the chicken embryo. Immunological aspects : Has one or more antigens in common with or closely resembling one or more present in the chlamydozoa and other miyagawanellae . Antisera against any of these two genera react with antigens from Miyagawanella lymphogranulomatis or the other miya- gawanellae thus far tested. One com- mon antigen has been isolated as a soluble fraction distinct from the bodies of the agent. Distinguished sharply from the other miyagawanellae by antitoxic neu- tralization of toxic factor or by neutra- lization of infections in mice with chicken antisera. Evidence exists that these two serological reactions are with dis- tinct specific antigens. Immunity in man or animals is probably poor in the absence of continuing apparent or inap- parent infection. Toxic factor: Infected yolk sac or yolk injected intravenously or intra- traperitoneally is rapidly fatal to mice. Produces characteristic lesions on the skin of normal guinea pigs. Pathogenicity : Pathogenic for man, apes, monkeys, guinea pigs, cotton rats, hamsters, mice, chicken embryos. In- apparent infections may occur with the agent harbored in the organs. Causes local genital lesions, septicemia, lymphad- enitis, meningitis, ophthalmitis and rarely pneumonitis in man. Tissue tropisms : In laboratory rodents this species is infective by the intra- nasal (pneumonitis), the intracerebral (meningitis) and the intradermal routes. Chemotherapy : Susceptible to rela- tively high concentrations of penicillin, to the sulfonamides and to some anti- mony compounds. Source : Most commonly the genital secretions of infected individual or the draining lymph nodes. Also occasionally in blood, spinal fluid and ocular secre- tions. Habitat : The etiological agent of lymphogranuloma venereum, lympho- granuloma inguinale, climatic bubo, es- thiomene and some forms of anorectal inflanunation . 2. Miyagawanella psittacii (Lillie) Aloshkovsky. (Rickettsia psittaci Lillie, Publ. Health Repts., 45, 1930, 773; Microbacterium multiforme psittacosis Levinthal,* 1st Cong, internat. de Mi- crobiol., 1, 1930, 523; Moshkovsky, Uspekhi Souremennoi Biologii (Russian) (Advances in Modern Biology), 19, 1945, 12; Ehrlichia psittaci Moshkovsky, ibid., 19.) From Psittaci, an order of birds. * This is the type species of the genus Microbacterium Levinthal which is invalid because of the earlier Microbacterium Orla-Jensen, 1919, see p. 370. FAMILY CHLAMYDOZOACEAE 1117 Coccoid bodies : As for Mhjagau-anclla lymphogranulornatis . Filterability : Partly filterable through Berkefeld N, Chamberland L and Q or Seitz EK filters. Cultivation : As for Miyagawanella lymphogrqnulomatis but grows readily in allantoic sac without adaptation. Immunological aspects: As for M. lymphogranulornatis but no soluble frac- tion yet demonstrated. Toxic factor: Infected yolk sac or yolk injected intravenously or intraperi- toneally is rapidly fatal to mice. Pathogenicity: Pathogenic for birds (particularly psittacine and finch species), man, monkeys, guinea pigs, pocket gophers, hamsters, white rats, kangaroo rat?, mice, rabbits and chicken embryos. Inapparent infections may occur with the agent harbored in the organs. Causes a highly fatal pneumoni- tis with septicemia in man. Tissue tropisms: Causes a septicemia. In man this species .shows predilection for the respiratory tract. In laboratorj^ rodents, it is infective by the intranasal, the intraperitoneal (peritonitis and septi- cemia), the intracerebral and the intra- venous routes. Chemotherapy: Susceptible to rela- tively high concentrations of penicillin. Some strains are susceptible to sulfon- amides. Source : Found in the organs and nasal secretions of infected birds and, from the latter, spreads to the plumage by preen- ingandothermethods. Plentiful in drop- pings or dust from infected cages. Rela- tively resistant under such conditions. Habitat: The etiological agent of psit- tacosis or parrot fever. Also of some cases of atypical pneumonia. 3. Miyagawanella ornithosis Rake, spec. nov. From Greek ornithos, bird. Coccoid bodies: As for Miyagawanella lymphogranulornatis . Cultivation : As for Miyagawanella psittaci. Immunological aspects : Has one or more antigens in common with, or closely resembling, one or more present in chla- mydozoa and other miyagawanellae as shown by a cross reaction in complement fixation tests. Sharply distinguished from other miyagawanellae by toxin- antitoxin neutralization or by neutrali- zation of infection in mice with chicken antisera. The latter test however sug- gests that the agent of meningopneumo- nitis (Francis and Alagill, Jour. Exp. Med., 68, 1938, 147) is this species rather than something distinct. Immunity in man or animals is probably poor except in the presence of continuing apparent or inapparent infections. Cross reac- tions suggest that Miyagawanella orni- thosis may be more closely related to Miyagawanella h/mphogrannlomatis than is M. psittaci. Toxic factor: As for Miyagawanella psittaci. Pathogenicity : Pathogenic for birds (especially non-psittacine species), man, ferrets, guinea pigs, hamsters, white rats, kangaroo rats, mice, rabbits and chicken embryos. Inapparent infec- tions may occur. Causes a moderately severe pneumonitis with septicemia in man. Tissue tropisms: Causes a septicemia. In birds and man shows a predilection for the lungs. In laboratory rodents, this species is infective by the intranasal, in- tracerebral, intravenous and (with rela- tively large inocula of most strains) in- traperitoneal routes. Chemotherapy : Susceptible to rela- tively large doses of penicillin. Not susceptible to sulfonamides. Source : Found in organs and nasal se- cretions of finches, pheasants (including domestic chickens), domesticated doves, fulmar petrels and other birds. Spreads from the secretions to plumage and drop- pings. Habitat: The etiological agent of orni- thosis (Meyer) and meningopneumonitis (Francis and Magill). 1118 MANUAL OF DETERMINATIVE BACTERIOLOGY 4. Miyagawanella pneumoniae Rake, spec. nov. Named for the disease, pneu- monia. Coccoid bodies: As for Miyagawanella lymphogranulomatis but slightlj' smaller, circa 200 millimicrons in diameter. Cultivation : As for Miyagawanella psittaci. Immunological aspects : As for Miya- gawanella psittaci. Distinct from Miya- gawanella ornithosis by the neutraliza- tion test with chicken antisera. Pathogenicity: Pathogenic for birds, man, cotton rats, hamsters, white rats, kangaroo rats, mice and chicken embryos. Causes a fatal pneumonitis in man. Tissue tropisms : As for Miyagawanella ornithosis. Chemotherapy : As for Miyagawanella ornithosis . Source : Occurs in lungs of infected hu- mans. Possibly originally of avian origin. Habitat: The etiological agent of one type of viral pneumonia. The type strain is the so-called strain S-F (Eaton, Beck and Pearson, Journ. Exp. Med., 73, 1941, 641). 5. Miyagawanella bronchopneumoniae Moshkovsky. (Aloshkovsky, Uspekhi Souremennoi Biologii, 19, 1945, 19; Ehrli- chia broncho pjieunwniae Moshkovsky, idem.) Named for the disease, broncho- pneumonia. Coccoid bodies: As for Miyagaivanella pneumoniae. Cultivation: As for Miyagawanelki lymphogranulomatis. Does not grow in the allantoic cavity of the chick. Immunological aspects : As for Miya- gawanella lymphogranulomatis but no soluble antigen has been demonstrated. Toxic factor : Heavily infected 3'olk sacs and yolk injected intravenously are very rapidly fatal to mice. Pathogenicity : Pathogenic for mice, hamsters and ferrets. Produces a mod- erately severe pneumonitis. Tissue tropisms: Shows a predileclion for the lungs. In mice, it is also infec- tive by the intravenous route. Chemotherapy : Susceptible to sulfon- amides and to relatively large doses of penicillin. Source : Found in lungs of certain stocks of the laboratory mouse. Habitat : The agent of mouse pneumo- nitis. Bronchopneumonie virus (Gon- nert. Cent. f. Bakt., I Abt., Orig., H7 , 1941, 151). 6. Miyagawanella felis Rake, spec. nov. From Latin Jelis, cat. Coccoid bodies : As for Miyagawanella lymphogranulomatis . Cultivation : As for Miyagawanella psittaci . Immunological aspects : As for Miya- gawanella psittaci but nothing known about inapparent infections in the natural host, the domestic cat. Toxic factor: Infected yolk sac or other membranes and yolk or other fluids, injected intravenously into mice or chicken embryos or intraperitoneally into mice are rapidly fatal. Pathogenicity : Pathogenic for cats, hamsters, mice and chicken embryos. Causes a fatal pneumonitis with acute conjunctivitis in cats. Tissue tropisms : Prediliction for lungs and conjunctivae. In laboratory ro- dents, this species is infective by the intranasal, intraperitoneal, intracerebral and intravenous routes. Chemotherapy : As for Miyagaivanella ornithosis. Source : Lungs of infected cats. Habitat : The etiological agent of one form of cat nasal catarrh, influenza or distemper (Baker, Science, 96, 1942, 475) and feline pneumonitis. 7. Miyagawanella louisianae Rake, spec. nov. Named for the State of Louisiana. Coccoid bodies : As for Miyagawanella psittaci. FAMILY CHLA MYDOZOACEA E 1119 Filterabiiity : Filters through Berke- feld N and Maudler 6, 7 and 9 filters. Cultivation : In the yolk sac of the chicken embryo. Immunological aspects : Indistinguish- able from other miyagawanellae by com- plement fixation tests with yolk sac an- tigens. Partly distinguished from Mi- yagawanella psittaci and M. ornithosis by active immunization in mice and guinea pigs. Pathogenicity : Pathogenic for man, guinea pigs, cotton rats, mice and chicken embryos. Slightly pathogenic for white rats, golden hamsters and deer mice. Macacus rhesus monkeys, rabbits, musk- rats and nutria are unaffected. Causes a highly fatal pneumonitis and septicemia in man. Tissue tropisms: Causes a septicemia. In man this species shows predilection for the respiratory tract. In laboratory rodents it is infective by the intranasal, intraperitoneal, intracerebral, intra- muscular and subcutaneous routes. Chemotherapy : As for Miyagawanella ornithosis. Source: Sputum and organs of in- fected persons. Habitat : The etiological agent of Louisiana pneumonia (Olson and Larson, U. S. Pub. Health Repts., 59, 1944, 1373), so-called Borg strain. 8. Miyagawanella illinii Rake, spec. nor. Xamed for the State of Illinois. Coccoid bodies : As for Miyagaicanella lymphogran ulomatis . Filterabiiity : Passes through Berke- feld N or W filters. Cultivation : In the yolk sac of chicken embryo. Immunological aspects: Distinguished from other miyagawanellae by neutrali- zation tests in mice with chicken anti- sera and partly from Miijagaivanella psit- taci, M. ornithosis and M. pneumonia by active inununization in mice. Pathogenicity : Pathogenic for man and white mice . Causes a highly fatal pneu- monitis in man. Tissue tropisms : Infective in mice by the intranasal, inti-aperitoneal, intra- cerebral and subcutaneous routes. Source : Lungs of infected persons. Habitat : The etiological agent called the Illinois virus (Zichis and Shaugh- nessy, Science, 102, 1945, 301). Genus III. Colesiota Rake, gen. nov. {Rickettsia Coles, 17th Rept. Direct. Vet. Serv. and An. Ind. Un. South Africa, 1931, 175.) Named for Prof. Coles who first studied these organisms. Pleomorphic cells which may be coccoid, triangular, rod-shaped or in the form of rings. Gram-negative. Intracytoplasmic habitat. The type species is Colesiota conjunctivae (Coles) Rake. 1. Colesiota conjunctivae (Coles) Rake, cotnb. nor. {Rickettsia conjuncti- vae Coles, 17th Rept. Direct. Vet. Serv. and An. Ind. Un. South Africa, 1931, 17.5; Chlamydozoon conjunclirac Moshkovsky, Uspeklii Souremennoi Biologii, 19, 1945, 19.) From M. L. conjunctira. the con- junctiva. Pleomorphic bodies : Average diameter 600 to 950 millimicrons. May be solid and coccoid, rod-shaped, or triangular, or in form of open rings or horse-shoes. No chains. Masses fi-equent. No cap- sule. Stains with ordinary aniline dyes but less intense]}' than bacteria. Gram- negative. Non-motile. Cultivation : Has never been culti- vated. Immunological aspects: Unknown. Pathogenicit}^ : Pathogenic for sheep, cattle and goats. Causes acute con- junctivitis and keratitis. Tissue tropisms : Affects only the con- junctiva and cornea. 1120 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : Found in scrapings of cornea or conjunctiva or in discharges from affected eyes. Etiological agent of infec- tious or specific ophthalmia in sheep, cattle and goats. 2. Colesiota conjunctivae-gallii (Coles) Rake, comb. nov. (Rickettsia conjuncti- vae-galli Coles, Onders. Jour. Vet. Sci. and Indust., I4, 1940, 469.) From con- junctiva and Latin galhis, hen. Pleomorphic bodies : Similar to Cole- siota conjunctivae. Stain purplish-red or blue with the Giemsa stain. Cultivation : Has never been culti- vated. Immunological aspects : Unknown. Pathogenicity : Pathogenic for the do- mestic fowl. Causes an acute conjunc- tivitis and keratitis. Tissue tropisms: As for Colesiota con- junctivae. Source : As for Colesiota conjuiictivae. Habitat : The etiological agent of one form of ocular roup in fowls. Appendix: The following are similar to or identical with the above: Rickettsia conjwictivae-bovis (Coles, South Afr. Vet. Med. Assoc, 7, 1936, 1) cannot be distinguished from Colesiota conjunctivae by any described character- istics. Rickettsia lestoquardi Donatien and Gayot. (Bull. Soc. Path. Exot., S5, 1942, 325.) Found in benign conjunctivitis in swine similar to that which occurs in ruminants. APPENDIX TO RICKETTSIALES 1121 * Appendix to Order Rickettsiales : The following are described species of intra- cytoplasmic and intranuclear parasites of Protozoa whose relationships to similar parasites of arthropods and vertebrates are not yet clear. All of the protozoon in- tracellular parasites are of larger size than typical members of Rickettsiales and some have been placed in genera (Cladothrix, Micrococcus) where the typical species do not live intracellularly. Genus A. Caryococcus Dangeard. (Compt. rend. Acad. Sci., Paris, 1S4, 1902, 1365.) Genus established for a bacterial parasite of the nucleus of Euglena; organisms rounded. The type species is Caryococcus hypertrophicus Dangeard. 1. Caryococcus hypertrophicus Dan- geard. (Compt. rend. Acad. Sci., Paris, 1S4, 1902, 1365.) Parasitic in the nucleus of a flagellate {Euglena deses). Occurs in the nucleus as an agglomera- tion of close-set, rounded corpuscles. The nucleus increases considerably in volimie, the chromatin is reduced to thin layers against the membrane, the interior of the nucleus is divided into irregular compartments by chromatic trabeculae. 2. Caryococcus cretus Kirby. (Univ. Calif. Publ. Zool., 49, 1944, 240.) Para- sitic in the nucleus of a flagellate {Tri- chonympha corbula) from the intestine of a termite (Procryptotermes sp.), Madagascar. Spherules 1 to 1.5 microns or more in diameter, in preparations appearing clear with usually a chromatic, sharply de- fined, crescentic structure peripherally or interiorly situated, sometimes with two such bodies or several chromatic granules ; parasitic in nucleus ; parasit- ized nucleus enlarged only moderately or not at all, chromatin altered but not greatly diminished in amount . 3. Caryococcus dilatator Kirby. (Univ. Calif. Publ. Zool., 49, 1944, 238.) Parasitic in the nucleus of flagellates {Trichonympha chattoni and other species of Trichonympha) from the intestine of termites {Glyptotermes iridipennis) , Aus- tralia, and other species. Spherules 0.5 micron or less in di- ameter, internally differentiated with stainable granule or stainable region pe- ripherally situated; parasitic in nucleus and nucleolus; nuc-^eus becomes greatly enlarged and the c hromatin mostly or entirely disappears. 4. Caryococcus invadens Kirby. (Univ. Calif. Publ. Zool., 49, 1944, 238.) Parasitic in the nucleus of a flagellate {Trichonympha peplophora) from the in- testine of a termite {Neotermes howa), Madagascar. Spherules 1 to 1 .5 microns in diameter, sometimes arranged in pairs, often in- ternally differentiated with stainable central or peripheral granules or stained areas ; parasitic in the nucleolus or endo- some and nucleus; parasitized nucleolus becoming greatly enlarged and crossed by trabeculae, eventually consumed; nu- cleus becoming moderately enlarged, but chromatin not disappearing. 5. Caryococcus nucleophagus Kirby. (Univ. Calif. Publ. Zool., 49, 1944, 236.) Parasitic in the nucleus of a flagellate {Trichonympha corbula) from the intes- tine of termites {Procryptotermes sp.), Madagascar, and three species of Kalo- tennes (s. 1.) from Madagascar. * Prepared by Prof. Harold Kirby, Jr. fornia, October, 1946. University of California, Berkeley, Call- 1122 MANUAL OF DETERMINATIVE BACTERIOLOGY Spherules with a diameter of about 0.5 terior or interior to the chromatin mass, micron, sometimes arranged in pairs, sometimes with a thicker, crescentic, stainable area of the periphery on one side; parasitic within the nucleus, ex- which may be diminished in amount, but does not disappear, nor is the parasitized nucleus appreciably enlarged. Genus B. Drepanospira Petschenko. (Arch. f. Protistenk., 22, 1911, 282.) Cell incurved in two spiral turns that arc not abrupt, one of the ends pointed, the other a little rounded, no flagella, movement helicoid by means of all the body, no cell division, endospores formed, regular spherical colonies formed by individuals at certain stages of development. The type species is Drepanospira vmlleri Petschenko. 1. Drepanospira miilleri Petschenko. (Mullerina paramecii Petschenko, Cent, f. Bakt., I Abt., Orig., 56, 1910, 90; Petschenko, Arch. f. Protistenk., 22, 1911, 252; see also Kirby, in Calkins and Sunmiers, Protozoa in Biological Re- search, 1941, 1036.) Parasitic in the cytoplasm of Paramecium caudatum. Developing from a group of curved rods in the cytoplasm to a large, ellipsoi- dal mass almost filling the body. Nu- clear portion occupying part of the cell. The author regards this genus as be- longing in the family Spirillaceae be- tween Spirosoma and Microspira. Genus C. Holospora Haffkine. (Ann. Inst. Past., 4, 1890, 151.) Genus established for bacterial parasites of the ciliate, Paramecium aurelia {■■ Paramecium caudatum ?). The type species is Holospora undulaia Haffkine. 1. Holospora undulata Haffkine. (Ann. Inst. Past., Paris, 4, 1890, 151.) In the nucronucleus of the ciliate Para- m,ecium aurelia (= P. caudatum?). Gradually tapered at ends; 1§, 2 and 2§ spiral turns; develops from a small, fusiform body which grows and divides transversely; brings about a great en- largement of the micronucleus, which becomes filled with the spirals (see Drepanospira mulleri Petschenko). 2. Holospora elegans Haffkine. (Haff- kine, Ann. Inst. Past., Paris, 4, 1890, 154; see also Kirby, in Calkins and Sum- iners. Protozoa in Biological Research, New York, 1941, 1035.) In the micro- nucleus of the ciliate, Paramecium aurelia (= P. caudatum ?). Vegetative stage fusiform; elongated, elliptical, nucleus-like body in some; divides equatorially, budding at one end; transformation into spore entails en- largement, clear space separating mem- brane at sides, spore pointed at ends. 3. Holospora obtusa Haffkin. (Haff- kine, Ann. Inst. Past., Paris, 4, 1890, 153; also see Fiveiskaja, Arch. f. Pro- tistenk., 65, 1929, 276.) In the macro- nucleus of the ciliate Paramecium au- relia {= P. caudatum ?). Spores not spiralled and both ends are rounded. Reproduction by fission, also by formation of a bud at one of the ex- tremities of the fusiform cell. Bodies with rounded ends 12 to 30 microns long; also spindle-shaped bodies with pointed APPENDIX TO RICKIOTTSIALES 1123 ends, 0.5 by 3 to 6 microns (Fiveiskaja, loc. cit.). The following species have been placed in genera belonging in the orders Chlamy- dobacteriales and Eubacteriales respec- tively : Cladothrix pelomyxae Vele3^ (Veley, Jour. Linn. Soc, Zool., 39, 1905, 375; see also Leiner, Arch. f. Protistenk., 4^', 1924, 282; Kirby, in Calkins and Sum- mers, Protozoa in Biological Research, New York, 1941, 1025; Hollande, Bull. Biol. France Belg., 79, 1945, 49.) In the cytoplasm of the rhizopod, Peloinyxa palustris and probably also other species of Pelomyxa. Rods, 1.5 to 22 microns or more in length, divided into several to many sections by transverse partitions, gen- erally aggregated in proximity to the nuclei, which may be thickly invested by close-set bacteria applied to the surface. Micrococcus batrochorum (sic) Yaki- nioff. (Arch. f. Protistenk., 7£, 1930, 137.) In the cytoplasm of the flagellate, Trichomonas bairachorum from the tree toad {Hyla arborea) . Also seen free in preparations of the intestinal contents of Hyla. Round, 1 to 1.5 microns in diameter, grouped generally in aggregates of ir- regular form, but also occur individually. XoTE : Further descriptions of bacte- rial and other parasites of Protozoa with bibliography will be found in Calkins and Summers, Protozoa in Biological Re- search, New York, 1941, 1009-1113 and in Kirby, Univ. of Calif. Pub. in Zoology, 53, 1946, 193-207. SUPPLEIVIENT NO. 2 ORDER VIRALES THE FILTERABLE VIRUSES Copyright 1948 by Francis O. Holmes Princeton, N. J. FILTERABLE VIRUSES* The so-called filterable viruses, today generally called merely viruses, are still of unknown affiliations so far as relationships to established groups of microorganisms are concerned. They are treated here as members of an order, consisting of 13 families, 32 genera and 248 species. Among viruses as we know them, there are three constituent groups that have come to be recognized, and to some extent named and classified, through the largely separate efforts of bacteriologists, animal pathologists, and plant pathologists. Taxonomic overlappmg of the three groups, viruses affecting bacteria, viruses having onlj'- animal hosts, and viruses invading higher plants, can hardly be justified as yet by available evidence. Nevertheless it has been shown that a single virus may multiply both in a plant host and in an insect vector. This seems to dispose of the thought that adaptation to a plant or animal enviromiient would necessarily pre- clude utilization of other sources of the materials needed for multiplication. For the present it seems feasible to continue \\dth the custom, tacitly accepted in the past, of classifying bacteriophages separately as one sub-group, \-iruses causing diseases in seed plants as a second sub-group, and those causing diseases in animals as a third sub-group. It should be recognized that this may prove to be only a temporary arrangement, necessary because we have no evidence to warrant taxonomic overlapping of the three groups and useful while we await critical investigations and possible development of a substitute plan capable of displaying natural relationships to better advantage. Eventually evidence may become available to show that some bacteriophages can infect higher plants or anmials and can mcrease in the new envirormient, or that viruses known to attack anmials or plants can similarly enlarge their host ranges. Or, there may be discoveries of common physical properties that would aid in formulating an interlocking classification, for which at present we lack any substantial basis. It is of especial significance now that the three fields be unified at least by a parallel development of nomenclature. Toward this end the pres- ent section of this supplement is directed. * Supplement No. 2 has been prepared by Francis O. Holmes, The Rockefeller In- stitute for Medical Research, Princeton, N. J., September, 1944. In this section, authorities for the names of plant hosts are in general as given by Gray's New Manual of Botany, 7th edition, and Bailey's Manual of Cultivated Plants, 1938 ; in each of these standard works will be found a list of abbreviations customarily used in botany in citing authorities for binomials. 1127 1128 MANUAL OF DETERMINATIVE BACTERIOLOGY ORDER VIRALES Breed, Murray and Kitchens. (Jour. Bact., J^^ , 1944, 421.) Viruses. Etiological agents of disease, typically of small size and capable of passing filters that retain bacteria, increasing only in the presence of living cells, giving rise to new strains by mutation, not arising de novo. A considerable num- ber of viruses have not been proved filterable; it is nevertheless customary to in- clude these viruses with those known to be filterable, because of similarities in other attributes and in the diseases induced. Some not known to be filterable are inoculable only by special techniques, as by grafting or by use of insect vectors, and suitable methods for testing their filterability have not been developed; more- over, it is not certain that so simple a criterion as size measured in terms of filter- ability will prove to be an adequate indicator of the limits of the natural group. Cause diseases of bacteria, plants and animals. Key to the suborders of order Virales. I. Infecting bacteria. Suborder I. Phagineae, p. 1128. II. Infecting higher plants. Suborder II. Phytophagineae p. 1145. III. Infecting animals (insects, mammals). Suborder III. Zoophagineae, p. 1225. Suborder I. Phagineae subordo novus. Viruses pathogenic in bacteria; bacteriophages. Containing at present only one family, the Phagaceae. FAMILY I. PHAGACEAE HOLMES. (Handb. Phytopath. Viruses,* 1939, 1.) Characters those of the suborder. There is a single genus. Genus I. Phagus Holmes. (Lac. cit., 1.) Characters those of the family. Generic name from Greek phagein, to eat. The type species is Phagus minimus Holmes. Note: Bacteriophagum d'Herelle (Cempt. rend. Soc. Biol., Paris, 81, 1918, 1161) a genus name applied in connection with early studies of bacteriophages, had as its type species Bacteriophagum intestinale d'Herelle, a bacteriophage that is not now identifiable or, more probably, a mixture of such unidentifiable bacteriophages, for filtrates containing it were said to be capable of killing outright a culture of bacteria (iftid., 1160). The genus name Bacierzop/iagfMm is, therefore, regarded as a nomen du- bium, if not also a nomen confusum; subsequently it was abandoned by its author, for reasons that are not clear, in favor of the genus name Protobios d'Herelle 1924 (Im- munity in natural infectious disease ; pagF 343 of authorized English edition by George H. Smith, Baltimore, Williams & Wilkins Co., 1924, 399 pp). Protobios protobios * Holmes, F. O., Handbook of Phytopathogenic Viruses, Burgess Publishing Com- pany, Minneapolis, Minn., 1939, 221 pp. FAMILY PHAGACEAE 1129 d'Herelle (loc. c?^, 345), presumably the type species of this genus, was not an ordinary virus but was said to be non-parasitic (i.e., free-living) in nature, was capable of re- ducing sulphur, and is not now identifiable. The genus name Protobios and the cor- responding binomial Protobios bacteriophagus d'Herelle are therefore regarded also as nomina diibia and are not used here. Bacteriophagus Thornberry (Phytopath., SI, 1941, 23) appears to represent a variant spelling of d'Herelle's earlier genus name; it was not accompanied by any indication of what recognizable single bacteriophage served as type and thus does not modif}' the standing of Bacteriophagum. Key to the species of genus Phagus. I. Dj'sentery-coli bacteriophages. A. Producing large plaques, 8 to 12 mm in diameter. 1. Particle size small, 8 to 12 millimicrons. 1. Phagus minivnis. 2. Particle size 15 to 20 millimicrons. 2. Phagus minor. B. Producing moderately large plaques, 2 to 6 mm in diameter, with distinct halo. 1. Particle size 20 to 30 millimicrons. 3. Phagus parvus. 4. Phagus primarius . 5. Phagus secundarius. G. Phagus dysenteriae. C. Plaques medium size, 1 to 3 mm in diameter, with distinct halo. 1. Particle size 25 to 40 millimicrons. 7. Phagus medius. 8. Phagus astrictus. D. Plaques small, 0.5 to 1.5 mm in diameter, with soft edge or narrow halo. 1. Particle size 30 to 45 millimicrons. 9. Phagus major. 10.' Phagus coli. 11. Phagus artus. E. Plaques very small, 0.1 to 1.2 mm in diameter, with sharp edges. 1. Particle size 50 to 75 millimicrons. 12. Phagus maximus. II. Bacteriophages attacking Agrobacterium tumefaciens Conn, Pseudomonas solanacearum Smith, Xanthomonas citri Dowson, Xanthomonas pruni Dowson, Erwinia carotovora Holland, Erwinia aroideae Holland, Bac- terium stewarti E. F. Smith. 1130 MANUAL OF DETERMINATIVE BACTERIOLOGY A. Specific for bacterial hosts named above. 13. Phagus tumoris. 14. Phagus solanacearum. 15. Phagus citri. 16. Phagus pruni. 17. Phagus defor?nans. 18. Phagus contumax. 19. Phagus maidis. III. Bacteriophages attacking Salmonella enteritidis Castellani and Chalmers. 20. Phagus enteritidis. 21. Phagus commutahilis. 22. Phagus tertius. 23. Phagus dubius. IV. Bacteriophage attacking Salmonella typhosa. 24. Phagus indicens. V. Bacteriophages attacking Bacillus megatherium DeBary, Bacillus my- coides Fliigge, and Rhizobium leguminosaruni Frank. A. Thermal inactivation at 75° C in 10 minutes in vitro. 1. Host may be freed from bacteriophage by heating at 80' C for 10 minutes. 25. Phagus teslabilis. 2. Host retains virus even when heated at 90* C for 10 minutes. 26. Phagus indomitus. B. Thermal inactivation at 60° C in 30 minutes. 27. Phagus subvertens. VI. Bacteriophages attacking streptococci. I 28. Phagus ineptus. 29. Phagus streptococci. 30. Phagus maculans. 31. Phagus lacerans. 32. Phagus tolerans. 33. Phagus michiganensis . VII. Bacteriophages attacking staphylococci . 34. Phagus fragilis. 35. Phagus intermedins. 36. Phagus caducus. 37. Phagus alpha. 38. Phagus beta. FAMILY PHAGACEAE 1131 39. Fhagus durabilia. 40. Phagus liber. VIII. Bacteriophages attacking vibrios. 41. Phagus cholerae. 42. Phagus celer. 43. Phagus effrenus. 44. Phagus lentus. IX. Bacteriophages attacking Corynebacterium diphthertae Lehmann and Neu- mann. 45. Phagus diphtheriae. 46. Phagus futilis. 1. Phagus minimus Holmes. (Handb. Phytopath. Viruses, 1939, 141.) From Latin minimus, least, in reference to size. Common name : Bacteriophage S13. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease : On plate cultures that are uniformly covered with confluent colonies of host organisms, this bacterio- phage produces large cleared plaques, 8 to 12 mm in diameter, with wide shelving edges. Serological relationships : Xo cross- neutralization reactions with bacterio- phages C13, C36, Do, D20, C18, D3, S8, C21, C16, andD6. Immunological relationships : Member of Resistance Group I. Other properties : Particle size 8 to 12 millimicrons. Xot affected by 26.3 per cent urea solution. Little or no inactiva- tion by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Lysis completely in- hibited by 0.25 per cent solution of sodium citrate. Literature : Burnet and Mclvie, Jour. Path, and Bact., 36, 1933, 299-306, 307- 318; 37, 1933, 179-184; Burnet et al., Austral. Jour. Exp. Biol, and Med. Sci., 15, 1937, 227-368. 2. Phagus minor H. {loc. cit., 141). From Latin minor, lesser. Common names : Bacteriophage C13, C8, and D44. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease: Large placjues, 8 to 12 mm in diameter, with wide shelving edges. Serological relationships : Cross reac- tions with bacteriophages C8 and D44 but not with bacteriophages S13, C36, D5, D20, D13, C18, D3, S8, C21, C16, D6. Immunological relationships : Member of Resistance Group I. Other properties : Particle size, 15 to 20 millimicrons. Completely inactivated by 1 : 25 ,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Specific soluble substance formed in lysed cultures blocks phage- antiphage reaction. Literature: Burnet, Jour. Path, and Bact., 36, 1933, 307-318; Brit. Jour. Exp. Path., i4, 1933, 100-108. 3. Phagus parvus H. {loc. cit., 142). From Latin parvus, small. Common names : Bacteriophage C36, S18, C38, M, and C37 of Burnet. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease : Moderately large plaques, 2 to 6 mm in diameter, with distinct halo. 1132 MANUAL OF DETERMINATIVE BACTERIOLOGY Serological relationships : Induces for- mation of antibody capable of neutraliz- ing bacteriophages S18, C38, M, and C37, but not bacteriophages S13, C13, D5, D20, D13, C18, D3, S8, C21, C16, or D6, which represent distinct serological groups . Immunological relationships : Member of Resistance Group I. Other properties : Particle size, 20 to 30 millimicrons. Completely inactivated by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Literature: Burnet, Jour. Path, and Bad., 36, 1933,307-318. 4. Phagus primarius H. {loc. cit., 143). From Latin primarius, chief or first. Common names : Bacteriophage Do, C51,C50, andD48. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease : Moderately large plaques, 2 to 6 mm in diameter, witli distinct halos. Serological relationships : Cross-neu- tralization reactions with bacteriophages C51, C50, and D48, but not with S13, C13, C36, D20, D13, C18, D3, S8, C21, C16, D6. Immunological relationships : Member of Resistance Group I. Other properties : Particle size, 20 to 30 millimicrons. Completely inactivated by 1 : 25 ,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Literature: Burnet, Jour. Path, and Bact., 36, 1933, 307-318. 5. Phagus secundarius H. {loc. cit., 143). From Latin secundarius, inferior or second. Common names : Bacteriophage D20 and G. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease : Moderately lai^e plaques, 2 to 6 mm in diameter, with dis- tinct halo. Serological relationships : No cross- neutralization reactions with bacterio- phages S13, C13, C36, D5, D13, CIS, D3, S8, C21, C16, or D6. Immunological relationships : Member of Resistance Group II. Other properties : Nearly all inacti- vated by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Particle size, 20 to 30 millimicrons. Literature: Burnet, Jour. Path, and Bact., 36, 1933, 307-318. 6. Phagus dysenteriae H. {loc. cit., 144). Fvoin lidtin dysenteria, dysentery. Common names : Bacteriophage D13, specific dysentery phage. Host : Shigella dysenteriae Castellani and Chalmers. Insusceptible species: Escherichia coli Castellani and Chalmers. Induced disease : Moderately large plaques, 2 to 6 mm in diameter, with distinct halo. Serological relationships : Antiserum to this strain is not known to be effective against any other strain of bacteriophage ; in particular, no cross reactions with bac- teriophages S13, C13, C36, D5, D20, C18, D3,S8, C21,C16,orD6. Immunological relationships : Member of Specific Dysentery Resistance Group. Other properties: Particle size, 20 to 30 millimicrons. Completely inactivated by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Literature : Burnet, Jour. Path, and Bact., 36, 1933, 307-318. 7. Phagus medius H. {loc. cit., 144). From Latin medius, moderate, in refer- ence to particle size. Common name : Bacteriophage C18, C35, C26, C47, or C34. Hosts : Escherichia coli Castellani and Chalmers; Shigella dysenteriae Castellani and Chalmers. FAMILY PHAGACEAE 1133 Induced disease : Medium size plaques, 1 to 3 mm in diameter, with distinct lialo. Serological relationships : Cross reac- tions with bacteriophages C35, C26, C47, and C34, but not with S13, C13, C36, D5, D20, D13, D3, S8, C21, C16, or D6. Immunological relationships : Member of Resistance Group II. Other properties: Particle size, 25 to 40 millimicrons. Literature: Burnet, Jour. Path, and Bact.,S6, 1933, 307-318. Serological relationships : No cross - neutralization reactions with bacterio- phages S13, C13, C36, Do, D20, D13, CIS, D3,C21,C16,orD6. Immunological relationships : Member of Resistance Group I. Other properties : Particle size, 30 to 45 millimicrons. Literature : Burnet, Jour. Path, and Bact., 36, 1933, 307-318; Brit. Jour. Exp. Path., 14, 1933, 100-108; Gough and Burnet, ibid., 38, 1934, 301-311. 8. Phagus astrictus H. {loc. cit., 145). From Latin astrictus, limited, in refer- ence to inability to Ij'se Escherichia coli Castellani and Chalmers. Common names : Bacteriophage D3 ; "smooth" dysentery phage. Host : Shigella dysenteriae Castellani and Chalmers. Insusceptible species : Escherichia coli Castellani and Chalmers. Induced disease : Medium size plaques, 1 to 3 mm in diameter, with distinct halo. Serological relationships: No cross- neutralization reactions with bacterio- phages S13, C13, C36, D5, D20, D13, C18, S8, C21,C16,orD6. Immunological relationships : Member of Smooth Dysentery Resistance Group. Other properties : Particle size, 25 to 40 millimicrons. Nearly all inactivated by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Literature: Burnet, Jour. Path, and Bact., 36, 1933,307-318. 10. Phagus coli H. {loc. cit., 146). From Latin colon, the colon. Conmion names : Bacteriophage C21 or C5; specific coli phage. Host : Escherichia coli Castellani and Chalmers. Insusceptible species : Shigella dysen- teriae Castellani and Chalmers. Induced disease : Small plaques, 0.5 to 1.5 mm in diameter, with soft edge or very narrow halo. Serological relationships : No cross- neutralization with bacteriophages S13, C13, C36, D5, D20, D13, C18, D3, S8, C16, or D6. Immunological relationships : Member of Specific Escherichia coli Resistance Group. Other properties : Particle size, 30 to 45 millimicrons. Completely inactivated by 1 : 25,000 methylene blue in 2 mm layer 20 cm from 100 candle-power light for 30 minutes. Literature: Burnet, Jour. Path, and Bact., 36,1933,307-318. 9. Phagus major H. {loc. cit., 146). From Latin major, greater, in reference to particle size. Common name : Bacteriophage S8, L, S28, C33,or S41. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease : Small plaques, 0.5 to 1.5 mm in diameter, with soft edge or narrow halo. 11. Phagus artus H. {loc. cit., 148). From Latin artus, narrow, in reference to limited host range. Common names : Bacteriophage D6, D33; smooth dysentery phage. Host : Shigella dysenteriae Castellani and Chalmers, smooth strains. Induced disease : Small plaques, 0.5 to 1.5 mm in diameter, with soft edge or very narrow halo. Serological relationships : Not neu- 1134 MANUAL OF DETERMINATIVE BACTERIOLOGY tralized by sera specific for bacterio- phages S13, C13, C36, D5, D20, D13, C18, D3, S8, C21,orC16. Immunological relationships : Member of Smooth Dysentery Resistance Group. Other properties: Particle size, 30 to 45 millimicrons. Literature: Burnet, Jour. Path, and Bact., 36, 1933, 307-318. 12. Phagus maximus H. {loc. cit., 147). From Latin maximus, greatest, in refer- ence to particle size. Common names : Bacteriophage C16, C4, C15, C20, C32, C46, D4, D12, D29, D53, H, J, K, and W. L. L. Hosts : Escherichia coli Castellani and Chalmers ; Shigella dysenteriae Castellani and Chalmers. Induced disease: Small plaques, 0.1 to 1.2 mm in diameter, with sharp edges. Serological relationships : No cross - neutralization reaction with bacterio- phages S13, C13, C36, D5, D20, D13, CIS, D3, S8, C21, D6, or staphylococcus bac- teriophage Au2. Agglutinated and inac- tivated by homologous, though not by other, antisera. For agglutination an original titer of 2 X lO^" or higher is re- quired; the reaction is visible to the unaided eye after 24 hours at 50° C and succeeds even after inactivation by heat (70 to 85° C for 30 minutes), formalde- hyde, or a photodynamic dye (pro- flavine). Immunological relationships : Member of Resistance Group II. Thermal inactivation : At or below 70° to 85° C for 30 minutes. Other properties : Particle size esti- mated by filtration as 50 to 75 millimi- crons, by centrifuging as 79 to 90 milli- microns, from photographs as 50 to 60 millimicrons. Rapidly inactivated by 26.3 per cent urea solution. Little or no inactivation by 1:25,000 methylene blue in 2 mm layer 20 cm from 100 candle- power light for 30 minutes. Lysis not inhibited by L5 per cent or weaker solu- tions of sodium citrate. Thermolabile specific soluble substance formed in lysed cultures blocks phage -antiphage reaction. Literature: Burnet, Brit. Jour. Exp. Path., 14, 1933, 93-100, 100-108, 302-308; Jour. Path, and Bact., 36, 1933, 307-318; 37, 1933, 179-184 ; Burnet and Lush, ibid., 40, 1935, 455-469; Burnet and McKie, ibid., 36, 1933, 299-306. 13. Phagus tximoris H. {loc. cit., 150). From Latin tumor, a swelling, in refer- ence to association of this bacteriophage with bacterial tumors. Common name : Agrobacierium tume- faciens bacteriophage. Host: Agrobacterium tumefaciens Conn, most strains. Insusceptible species : Some strains of Agrobacterium tumefaciens, Bacterium stewarti E. F. Smith, Erwinia atroseptica Bergey et al., E. carotovora Holland, Pseudomonas tabaci Stapp, Xanthomonas beticola Burkholder, X. campestris Dow- son, X. citri Dowson, X. phaseoli Dowson X. pruni Dowson and X. vesicatoria Dowson. Geographical distribution : United States, Russia. Induced disease : Plaques 2 to 6 mm in diameter in 4 to 6 hours, edges of plaques spotted, moth-eaten in appearance until 40 hours after seeding; enlargement then stops and the edges of the plaques become smooth, double-ringed. Infection of plants by Agrobacterium tumefaciens is progressively inhibited by increasing amounts of bacteriophage in inoculum. Thermal inactivation: At 95° C in 10 minutes (another report says 70° C, time not recorded). Other properties : Resists dilution to 1 : 10" ; storage at 5° C for over 25 months ; prompt, though not gradual, drying; 1 per cent hydrogen peroxide for 72 hours ; 95 per cent ethyl alcohol for 1 hour; 70 per cent ethyl alcohol for 6 hours ; 2| per cent phenol for 1 hour; 1 :3000 nitric acid for 1 hour; N/64 sodium hydroxide for 1 hour. Literature: Israilsky, Cent. f. Bakt., FAMILY PHAGACEAE 1135 II Abt., 67, 1926, 236-242; 71, 1927, 302- 311 ; 79, 1929, 354-370; Kent, Phytopath., 27, 1937, 871-902; Muncie and Patel, Phytopath., 20, 1930, 289-305. 14. Phagus solanacearum H. {loc. cit., 148). From name of host. Common name : Pseudomonas solanace- arum bacteriophage. Host : Pseudomonas solanacearum Smith. Geographical distribution : Formosa (Taiwan). Induced disease : Medium size plaques on plate cultures of Pseudomonas sola- nacearum. Serological relationships : When in- jected into rabbits, this bacteriophage stimulates the production of a specific precipitating antibody not giving cross reactions with anti-bacterial antibodies. Antiphagic serum inactivated at 90° C in 10 minutes. Thermal inactivation: At 63° C in 10 minutes (61° C in 30 minutes; 66° C in about 1 minute). Other properties : Optimum tempera- ture for increase, 34° C. Literature: Matsumoto and Okabe, Jour. Plant Prot., 22, 1935, 15-20; Jour. Soc. Trop. Agr., 7, 1935, 130-139; .9, 1937, 205-213. 15. Phagus citri H. {loc. cit., 149). From name of host. Common name : Xanthomonas citri bacteriophage. Host : Xanthomonas citri Dowson, the citrus canker organism. Geographical distribution : Formosa (Taiwan). Induced disease: Lysis. This bac- teriophage has been isolated from soil under diseased trees, and once from infected leaves. It may play a role in the destruction of the citrus canker organism in the soil. Other properties : Optimum tempera- ture for increase, 30° C. Literature : Matsumoto and Okabe, Agriculture and Horticulture, 12, 1937, 2055-2059. 16. Phagus pruni H. (loc. cit., 151). From name of host. Common name : Xanthomonas pruni bacteriophage. Host : Xanthomonas pruni Dowson. Geographical distribution : United States (from soil beneath infected peach trees). Induced disease : Lysis in broth cul- tures; plaques on agar cultures, but characteristics of plaques not described. Other properties : Estimated diameter 11 millimicrons in broth. Resists dilu- tion to 1 :10® or more. Literature: Anderson, Phytopath., 18, 1928, 144; Thornberry, ibid., 25, 1935, 938-946. 17. Phagus deformans H. {loc. cit., 151 ) . From Latin deformare, to disfigure, in reference to malformation of infected host cells. Common name : Erwinia carotovora bacteriophage . Host : Erxoinia carotovora Holland. Insusceptible species : Agrobacterium tumefaciens Conn, e.xcept in some early tests with possibly mixed bacteriophages ; Erwinia amylovora Winslow et al., E. melonis Holland, Salmonella pullorum Bergery et al., *S. gallinarum Bergey et al.. Shigella dysenteriae Castellani and Chalmers, Xanthomonas pruni Dowson. Geographical distribution : United States (Michigan). Induced disease: In Erwinia caroto- vora, cells reduced in motility, agglu- tinated, malformed, some elongated, others swollen, bulged at one end, bulged in middle, or enlarged and spherical. Other properties : Resists dilution to 1:10% and storage in sterile medium at room temperature for 5| months. Literature : Coons and Kotila, Phyto- path., 15, 1925, 357-370; Mallmann and Hemstreet, Jour. Agr. Res., 28, 1924, 599-602. 1136 MANUAL OF DETERMINATIVE BACTERIOLOGY 18. Phagus contumax spec. TOov. From Latin contumax, refractory, in reference to ability of this bacteriophage to with- stand heating sufficient to destroy accom- panying host cells. Common name : Erwinia aroideae bac- teriophage. Host : Erwinia aroideae Holland. Insusceptible species : Agrohacterium iumefaciens Conn, Bacterium formosanum Okabe, Erwinia carotovora Holland, Pseudomonas andropogoni Stapp, P. solanacearum Smith, P. tomato Burk- holder, Xanthomonas campestris Dowson, X. citri Dowson, X. malvacearum Dow- son, X. nakatae Dowson, X. phaseoli Dowson, X. ricinicola Dowson. Geographical distribution : Formosa (Taiwan). Induced disease : Very small plaques, 0.1 to 1.0 mm (mostly less than 0.5 mm) in diameter. Thermal inactivation : Resists heating at 60° C for 30 minutes without appreci- able loss of titer, but host organism is killed by this treatment. Other properties : Optimum tempera- ture for increase, about 25° C. This bac- teriophage may be prepared by heating centrifuged cultures at 60° C for 30 min- utes as efficiently as by filtration to re- move bacteria. Literature: Matsumoto, Trans. Nat. Hist. Soc. Formosa, 29, 1939, 317-338; SO, 1940, 89-98; 31, 1941, 145-154; Mat- sumoto and Sawada, ibid., 28, 1938, 247-256. 19. Phagus maidis H. (loc. cit., 152). From New Latin 77iais, corn (maize), host of Bacterium stewarti. Common name : Bacterium stewarti bacteriophage; Phytomonas stewarti bac- teriophage; Aplanobacter stewarti bac- teriophage. Host: Bacterium stewarti E. F. Smith (= Pseudomonas stewarti E. F. Smith, Phytomonas stewarti Bergey et al. and Aplanobacter stewarti McCulloch). Geographical distribution : United States. Induced disease : In Bacterium ste- warti, variation or loss of yellow color, change of viscosity of growth, reduction or loss of virulence. Infection of corn plants by seed-borne Bacterium stewarti is much reduced by treating seeds with this bacteriophage before they are planted. Thermal inactivation : Above 65° C in 30 minutes. Other properties : Infective in dilu- tions to 10"^. Soon lost from cultures maintained at pH 3.85 to 4.00, or on Ivanoff's medium, which contains oxidiz- ing compounds. Literature: Thomas, Phytopath., 25, 1935, 371-372; Science, 88, 1938, 56-57; Phytopath., 30, 1940, 602-611. 20. Phagus enteritidis H. {loc. cit., 153). From name of host. Conunon names : Salmonella enteritidis bacteriophage 1, 12, or 33; Group A bac- teriophages. Hosts : Salmonella enteritidis Castellani and Chalmers, *S'. gallinarum Bergey ct al., Shigella diisetiicriae Castellani and Chalmers. Induced disease : Plaques of medium size, usually with surrounding translu- cent halo. Immunological relationships : Member of Resistance Group A; host individuals that have acquired resistance to this bacteriophage are resistant to lines 12 and 33, but susceptible to Salmonella enteritidis bacteriophages 8, 20, and 11, as well as to other strains of Resistance Groups B, C, and D. Literature: Burnet, Jour. Path, and Bact., 32, 1929, 15-42. 21. Phagus commutabilis H. (loc. cit., 153). From Latin commutabilis, vari- able, in reference to differences within Resistance Group B, typified by this bac- teriophage. Common names : Salmonella enteritidis bacteriophage 8, 18, 28, 31, 34, 38; Group B bacteriophages. Hosts : Salmonella enteritidis Castellani FAMILY PHAGACEAE 1137 and Chalmers, Shigella dyscnteriae Cas- tellani and Chalmers, Shigella gallinarinn Weldin, Salmonella typhosa White. Induced disease : Small plaques with sharp edges, or moderately large plaques with characteristic halo. Immunological relationships : Member of Resistance Group B ; host individuals that have acquired resistance to this bac- teriophage are resistant to lines 18, 28, 31, 34, and 38, but susceptible to Sal- monella enteritidis bacteriophages 1, 20, and 11, as well as to other strains of Re- sistance Groups A, C, and D. Literature: Burnet, Jour. Path, and Bact., 3£, 1929, 15-42. 22. Phagus tertius H. {loc. cit., 154). From Latin tertius, third, in reference to the third Resistance Group of Salmonella enteritidis bacteriophages, Group C, typi- fied by this bacteriophage. Common names : Salmonella enteritidis bacteriophage 20, 25, 32, 35; Group C bacteriophages. Hosts : Salmonella enteritidis Castellani and Chalmers, S. gallinarum Bergcy et al., Shigella dysenteriae Castellani and Chalmers. Induced disease : Plaques of small size, with sharp edges. Immunological relationships : Member of Resistance Group C. Host individuals that have acquired resistance to this bac- teriophage are resistant to lines 25, 35, and 32, but susceptible to Salmonella enteritidis bacteriophages of Resistance Groups A, B, and D. Literature: Burnet, Jour. Path, and Bact., 3^, 1929,15-42. 23. Phagus dubius H. (loc. cit., 155). From Latin dubius, doubtful, in reference to uncertainty of distinction between Resistance Groups C and D. Common names : Salmonella enteritidis bacteriophage 11, 13; Group D bacterio- phages . Hosts : Salmonella enteritidis Castellani and Chalmers, Shigella dysenteriae Cas- tellani and Chalmers, Shigella gallinarum Weldin. Induced disease : Very large plaques, up to 8 mm in diameter on 1.2 per cent agar. Immunological relationships : Member of Resistance Group D. Host individ- uals that have acquired resistance to this bacteriophage are resistant to line 13, but susceptible to Salmonella enteritidis bac- teriophages of Resistance Groups A, B, and C. Literature: Burnet, Jour. Path, and Bact., 32, 1929, 15-42. 24. Phagus indicens spec. not'. From Latin indicere, to disclose or indicate, in reference to diagnostic use of this bac- teriophage in identifying V forms of the typhoid bacillus. Common name : Phage Q151. Host: Salmonella typhosa White ( = Bacillus typhosus Zopf). Insusceptible species : W forms of the typhoid organism and various Salmonella species. Geographical distribution : Canada. Induced disease : In Salmonella typhosa, small plaque formation (lysis) and com- plete inhibition of growth in cultures of the V form (bearing Vi antigen ; resisting O agglutination) and no lysis or restrain- ing effect on growth of the W form (lack- ing Vi antigen; agglutinated by O anti- serum). In the presence of the virus, mixed cultures are ciuickly transformed since only W variants can increase . Pure V cultures can be identified by the test for their complete inhibition; this in- hibition is regularly followed by second- ary growth representing the pure W form of the host, a readily formed variant. Filterability : Passes Seitz EK filter. Other properties : Filtrates active in dilutions to 10"^ or lO"". Literature: Craigie, Jour. Bact., 31, 1936, 56 (Abst.); Craigie and Brandon, Jour. Path, and Bact., 43, 1936, 233-248, 249-260. 1138 MANUAL OF DETERMINATIVE BACTERIOLOGY 26. Phagus testabilisH. {loc.cit., 155). From Latin testabilis, able to bear wit- ness, in reference to evidence that this bacteriophage has given, by virtue of its easy destruction when heated in spores, against the hypothesis of frequent spon- taneous origin of bacteriophage from the bacterial host. Common name : Bacillus megaihcrivm bacteriophage. Host : Bacillus megatherium De Bary. Geographical distribution : United States. Induced disease : Plaques 0.5 mm or less in diameter, with surrounding translu- cent zone. Thermal inactivation : In vitro, at 75° C in 10 minutes. Spores from infected cultures, after being heated for 10 min- utes at 80° C, regularly give rise to sub- cultures that do not show the presence of this bacteriophage spontaneously during subsequent growth but that are sus- ceptible to lysis if the bacteriophage is again introduced. Literature: Adant, Compt. rend. Soc. Biol., Paris, 99, 1928, 1246; Cowles, Jour. Bact., £0, 1930, 15-23. 26. Phagus indomitus H. {loc. cit., 156). From Latin indomitus, unre- strained, in reference to the ability of this bacteriophage to increase after heat treat- ment of infected spores. Common name : Bacillus mycoides bac- teriophage. Host : Bacillus mycoides Fliigge, some strains. Insusceptible species : Bacillus cereus Frankland and Frankland, B. stibiilis Cohn emend. Prazmowski, B. mega- therium De Bary, B. anthracis Cohn emend. Koch. Some strains of B. my- coides. Geographical distribution. United States. Induced disease: Large plaques, with some secondary growth of host organism. Thermal inactivation : In vitro, at 75° C in 10 minutes. Spores from infected cul- tures, heated at 90° C for 10 minutes give no bacteriophage on grinding, but lytic cultures when grown. liiterature : Lewis and Worley, Jour. Bact., 32, 1936, 195-198. 27. Phagus subvertens H. {loc. cit., 156). From Latin subvertere, to subvert, in reference to suspected action of this bacteriophage in causing running-out of alfalfa fields through destruction of nodule organisms. Common name : Rhizobium legumino- sarum bacteriophage. Host : Rhizobium leguminosariim Frank. It has been shown that this bacteriophage is unable to increase in clover roots without the nodule-forming organism, R. Icguminosarum, and that the bacteriophage plays no obviously essential role in nodule formation. Induced disease : Very small plaques, with edges not sharply defined. Thermal inactivation: At 60° C in 30 minutes. Other properties: Not inactivated by drying for 2 months. Literature: Gerretsen et al.. Cent. f. Bakt., II Abt., 60, 1923, 311-316; Grijns, ibid., 71, 1927, 248-251; Hitchner, Jour. Bact., 19, 1930, 191-201; Vandecaveye and Katznelson, Jour. Bact., 31, 1936, 465-477. 28. Phagus ineptus H. (loc. cit., 157). From Latin ineptus, unsuitable, in refer- ence to inability of this bacteriophage to adapt itself to lysis of strain RW of its host. Common name : Streptococcus bacterio- phage R. Host : Streptococcus cremoris Orla-Jen- sen, strain R. Insusceptible species : Streptococcus cremoris, strain RW. Geographical distribution : New Zea- land. Induced disease : Plaques 0.25 to 0.6 mm in diameter. Serological relationships : Antisera spe- cific for streptococcus bacteriophage RW FAMILY PHAGACEAE 1139 and its strain RWl are ineffective in neu- tralizing this bacteriophage. Immunological relationships : Cultures of host-strain R, after exposure to this bacteriophage, furnish subcultures only partly resistant to this bacteriophage and completely susceptible to streptococcus bacteriophage RW and its substrain RWl . Literature : Whitehead and Hunter, Jour. Path, and Bact., U, 1937, 337-347. 29. Phagus streptococci H. {loc. cit., 158). From generic name of host. Common name : Streptococcus bacterio- phage RW. Host : Streptococcus cremoris Orla-Jen- sen, strain RW. Geographical distribution : New Zea- land. Induced disease : Plaques 0.25 to 0.6 mm in diameter. Thermal inactivation : At 70° to 75° C, time not recorded, probably 30 minutes (pH6.0). Literature : Wliitehead and Hunter, Jour. Path, and Bact., U, 1937, 337-347. Strains : One variant has been described and distinguished from the type variety, typicus H. {loc. cit., 158) : 29a. Phagus streptococci var. virilis H. {loc. cit., 158). From Latin virilis, vig- orous. Common name: Strain RWl of streptococcus bacteriophage RW. Dif- fering from the type variety in being able to increase at the expense of strain RWl of Streptococcus cremoris (Whitehead and Hunter, Jour. Path, and Bact., 44, 1937, 337-347). 30. Phagus maculans spec. nov. From Latin maculare, to speckle, in reference to tiny plaques produced by this bac- teriophage . Common name : Streptococcus bac- teriophage A. Hosts : Streptococcus 646, 751, 775. Geographical distribution : United States (Massachusetts ) . Induced disease : Plaques exceedingly minute, scarcely visible to the unaided eye. Serological relationships : Specific anti- sera neutralize but there is no cross reac- tion with respect to streptococcus bac- teriophage B, C, or D. Thermal inactivation : At 60° C in 1 hour. Other properties : Withstands storage at about 5° C for at least 145 days with but little loss of virulence. Literature : Evans, Science, 80, 1934, 40-41; U.S.P.H.S., Public Health Re- ports, 49, 1934, 1386-1401. 31. Phagus lacerans spec. nov. From Latin lacerare, to tear, in reference to ragged edges of plaques produced by this bacteriophage. Common name : Streptococcus bac- teriophage B. Hosts : Streptococcus 563,639 : Strepto- coccus mucosiis Howard and Perkins. Insusceptible species : Streptococcus erysipelatos Rosenbach. Geographical distribution : United States (Wisconsin). Induced disease : Medium size plaques, the largest about 3 mm in diameter, edges ragged, centers clean. Serological relationships : Specific neu- tralization, but no cross reactions with streptococcus bacteriophages A, C, andD. Thermal inactivation : At 60° C in 1 hour. Other properties : Withstands storage at about 5° C for at least 261 days. Literature : Clark and Clark, Jour. Bact., U, 1926, 89; Proc. Soc. Exp. Biol, and Med., U, 1927, 635-639; Colvin, Jour. Inf. Dis., 51, 1932, 17-29; Evans, U.S.P.H.S., Public Health Reports, 49, 1934, 1386-1401; Jour. Bact., 39, 1940, 597-604; Shwartzman, Jour. Exp. Med., 46, 1927, 497-509. 32. Phagus tolerans spec. nov. From Latin tolerans, tolerating, in reference to the unusual ability of this streptococcus 1140 MANUAL OF DETERMINATIVE BACTERIOLOGY bacteriophage to remain viable under certain adverse conditions. Common name : Streptococcus bacterio- phage C. Hosts : Streptococcus 646, 594, 756, 806. Geographical distribution : United States (Ohio, Massachusetts, Connecti- cut). Induced disease: Small plaques, the largest about 1.0 nun in diameter. Serological relationships : Specific neu- tralization, but no cross reactions with streptococcus bacteriophages A, B, and D. Thermal inactivation : At 63° to 65° C in 1 hour. Other properties : Withstands storage in 1 :200 phenol at about 5° C for at least 261 days ; equally resistant to storage in 1 : 10,000 sodium ethyl mercurithiosalicy- late (merthiolate), or to storage without preservatives. Literature: Evans, U. S. Pub. Health Ser., Public Health Reports, 49, 1934, 1386-1401. 33. Phagus michiganensis spec. nov. From name of state, Michigan, where this bacteriophage was first isolated. Common name : Streptococcus bacterio- phage D. Host : Streptococcus 693. Geographical distribution: United States (Michigan). Induced disease: Small plaques, about 0.75 mm in diameter, edges clear-cut, centers clean. Serological relationships : Specific neu- tralization, but no cross neutralization with streptococcus bacteriophages A, B, and C. Thermal inactivation : At 60° to 63° C in 1 hour. Other properties : Withstands storage at about 5°C for at least 261 days. Literature: Evans, V. S. Pub. Health Ser., Public Health Reports, 49, 1934, 1386-1401. 34. Phagus fragilis, H. {loc. cit., 159). From haiin fragilis, fragile, in reference to easy destruction of this bacteriophage bj'^ light and by concentrated urea solu- tions. Common names : Staphylococcus bac- teriophage Au2, Au3, Au4, or D, perhaps bacteriophage H of Gratia. Hosts : Staphylococcus aureus Rosen- bach and Staphylococcus albus Rosenbach. Geographical distribution : United States . Induced disease : Small plaques, 0.2 to 1.0 mm in diameter, with sharp edges. Serological relationships : Cross-neu- tralization reactions with staphylococcus bacteriophages Aul, Au3, Au4, and D, but not with staphylococcus bacterio- phages Au21, Aul2, A, B, C, or bacterio- phage CI 6. Thermal inactivation : At about 57° C in 30 minutes. Other properties : Particle diameter 50 to 75 millimicrons. Readily inactivated photodynamically. Completely inacti- vated by 27 per cent urea solution in 1 hour at 37° C. Lysis not inhibited even by 1.5 per cent sodium citrate in agar medium. Literature : Burnet and Lush, Jour. Path, and Bad., 40, 1935, 455-469 ; Burnet and McKie, Austral. Jour. Exp. Biol, and Med. Sci., 6, 1929, 21-31 ; Fisk, Jour. Inf. Dis., 71, 1942, 153-160. 35. Phagus intermedius H. (loc. cit., 160). From Latin intermedius, inter- mediate, in reference to position of this bacteriophage between staphylococcus bacteriophages that multiply readily in broth cultures of host organisms and those that do not. Common name : Staphylococcus bac- teriophage Au21. Host : Staphylococcus aureus Rosen- bach. Geographical distribution : Australia. Induced disease : Small plaques, 0.1 to 0.3 mm in diameter, with sharp edges. Serological relationships : Specific neu- tralization reaction but no cross-neutrali- zation reaction with staphylococcus bac- teriophages Au2 or Aul2. FAMILY PHAGACEAE 1141 Other properties : Not readily inacti- vated photodynamically ; completely in- activated by 27 per cent urea solution in 1 hour at 37° C; lysis inhibited by 1 per cent sodium citrate in agar medium but not by 0.5 per cent or lower concentra- tions. Literature: Burnet and Lush, Jour. Path, and Bact., 40, 1935, 455-469. 36. Phagus caducus H. {loc. cit., 160). From Latin caducus, perishable, in refer- ence to the easy destruction of this bac- teriophage by concentrated urea solu- tions . Common name: Staphylococcus bac- teriophage Aul2. Host : Staphylococcus aureus Rosen- bach. Geographical distribution: Australia. Induced disease : Small plaques, 0.2 to 0.5 mm in diameter, with sharp edges. Serological relationships : Cross-neu- tralization reactions with staphylococcus bacteriophages Aull and Aul3, but not with staphylococcus bacteriophages Au2, Au21, A, and C. Antiserum to staphylo- coccus bacteriophage B gives no neutrali- zation of Aul2, though the reciprocal re- action occurs to 1 :200 dilution. Other properties: Not readily inacti- vated photodynamically; completely in- activated by 27 per cent urea solution in 1 hour at 37° C ; lysis inhibited by as little as 0.25 per cent sodium citrate in agar. Literature: Burnet and Lush, Jour. Path, and Bact., 40, 1935, 455-469. 37. Phagus alpha H. (loc. cit., 161). From Greek equivalent of common name. Common name : Staphylococcus bac- teriophage A. Host : Staphylococcus albus Rosenbach. Geographical distribution: Australia. Induced disease : Plaques of medium size, 1.5 to 2.5 mm in diameter, with hazy peripherJ^ Serological relationships : Specific neu- tralization reaction, but no cross-neu- tralization reactions with staphylococcus bacteriophages Au2, B, or C. Immunological relationships : Colonies of Staphylococcus albus appearing after lysis with this bacteriophage are resistant to staphylococcus bacteriophages B, C, and D. Thermal inactivation : .\t 68° to 70° C in 30 minutes. Other properties : Not readily inacti- vated photodynamically; not completely inactivated by 27 per cent urea solution in 1 hour at 37° C ; lysis not inhibited even by 1.5 per cent sodium citrate in agar. Literature: Burnet and Lush, Jour. Path, and Bact., 40, 1935, 455-469 ; Burnet and ^IcKie, Austral. Jour. Exp. Biol, and :\Ied. Sci.,6, 1929, 21-31. 38. Phagus beta H. (loc. cit., 162). From Greek equivalent of common name. Common name : Staphylococcus bac- teriophage B. Host : Staphylococcus albus Rosenbach. Geographical distribution: Australia. Induced disease : Plaques of medium size, 0.7 to 1.5 mm in diameter, with sharp edges. Serological relationships : Specific neu- tralization reaction, but no cross-neu- tralization reaction with respect to staphylococcus bacteriophages Au2, Aul2, A, or C, except that antiserum made with Aul2 neutralizes this bacteriophage in low dilutions (See Phagus caducus). Immunological relationships : Colonies appearing after lysis of Staphylococcus al- bus with this bacteriophage furnish organ- isms susceptible to staphylococcus bac- teriophages A and D. Thermal inactivation : At 63° to 65° C in 10 minutes. Other properties: Readily inactivated photodynamically; completely inacti- vated by 27 per cent urea solution in 1 hour at 37° C ; lysis not inhibited even by 1.5 per cent sodium citrate in agar medium. Literature: Burnet and Lush, Jour. Path, and Bact., 40, 1935, 455-469 ; Burnet 1142 MANUAL OF DETERMINATIVE BACTERIOLOGY and McKie, Austral. Jour. Exp. Biol, and Med. Sci., 6, 1929,21-31. 39. Phagus durabilis H. (loc. cit., 162). From Latin durabilis, lasting, in refer- ence to the stability of this bacteriophage in concentrated urea solution and other unfavorable media. Common name : Staphylococcus bac- teriophage C. Host : Staphylococcus albus Rosenbach. Geographical distribution : Australia. Induced disease : Plaques 2.0 to 3.0 mm in diameter. Vitreous change in pe- ripheral zone. Serological relationships : Cross-neu- tralization reaction with staphylococcus bacteriophage C, and less strongly with B, but not with Au2 or A. Immunological relationships : Colonies of Staphylococcus albvs appearing after lysis with this bacteriophage furnish or- ganisms resistant to it but susceptible to staphylococcus bacteriophages A, B, and D. Thermal inactivation : At 61° to 63° C in 30 minutes. Other properties : Not readily inacti- vated photodynamically ; not completely inactivated by 27 per cent urea solution in 1 hour at 37° C ; lysis not inhibited even by 1.5 per cent sodium citrate in agar medium. Literature: Burnet and Lush, Jour. Path, and Bad., 40, 1935, 455-469 ; Burnet and McKie, Austral. Jour. Exp. Biol, and Med. Sci., 6, 1929, 21-31 ; Rakieten et al., Jour. Bad., 32, 1936, 505-518. 40. Phagus liber H. {loc. cit., 163). From Latin liber, independent, in refer- ence to demonstrated independence of this virus, its bacterial host, and its dip- terous superhost, in respect to origin. Common name : Staphylococcus muscae bacteriophage. Host : Staphylococcus muscae Glaser. Geographical distribution: United States. Induced disease : Lysis in broth cul- tures ; plaques in agar cultures, but char- acteristics of plaques not recorded. Thermal inactivation : At a little above 50° C in 5 minutes. Other properties : A characteristic nucleoprotein has been isolated from lysed staphylococci. Sedimentation con- stant, 650 X 10-1' cm dyne-i sec.-i, corresponding to a molecular weight of about 300,000,000. Denatured at acidi- t ies beyond pH 5.0. Digested by chymo- trypsin, not by trypsin. Apparent den- sity, about 1.20. Diffusion coefficient, varying with dilution. Literature : Glaser, Amer. Jour. Hy- giene, 27, 1938, 311-315; Northrop, Jour. Gen. Physiol., 21, 1938, 335-366; Shope, Jour. Exp. Med., 45, 1927, 1037-1044; Wyckoff, Jour. Gen. Physiol., 21, 1938, 367-373. 41. Phagus cholerae H. {loc. cit., 164). From former name of host. Common name : Vibrio comma bacterio- phage. Host: Vibrio comma Winslow et al. (formerly V. cholerae Neisser) ; Indian strains usually carry this bacteriophage, but Chinese and Japanese strains lack it, are susceptible, and upon inoculation be- come lysogenic. Geographical distribution : India. Induced disease : In both R and S forms of Vibrio comma, no plaques on ordinary agar plates, but vibrios become l3^sogenic. Egg-white in 1:25 dilution enhances activity enough to allow visible lysis, occasional plaques, or stippling at the site of inoculation. Immunological relationships : Vibrio comma organisms that have been infected with this bacteriophage and are resistant to its further action are still susceptible to cholera bacteriophages A, C, and D. Literature: White, Jour. Path, and Bact.,.^.^, 1937,276-278. 42. Phagus celer H. {loc. cit., 164). From Latin celer, quick, in reference to relatively quick action of this bacterio- phage. FAMILY PHAGACEAE 1143 Common name: Cholera bacterio- phage A. Host: Vibrio comma Winslow et al., smooth types, except non-agglutinable vibrios. Geographical distribution: India. Induced disease : Lj^sis in 2 hours, fol- lowed by abundant secondary growth. Onlj' smooth elements of the culture are attacked. Serological relationships : Antigenicallj' distinct from cholera bacteriophage C. Immunological relationships : Second- ary growth resistant to this virus, but susceptible to cholera bacteriophages C and D. Other properties: Selective]}^ inacti- vated by specific polysaccharide of smooth strains, not by a lipoid emulsion that is effective against cholera bacterio- phage C. Active in dilution of 1:10" or 1:10'". Multiplication rate, n X 10^ in 2 hours. Literature: Asheshov et al., Indian Jour. Med. Res., 20, 1933, 1127-1157; White, Jour. Path, and Bact., 43, 1936, 591-593. 43. Phagus effrenus H. {loc. cit., 165). From Latin effrenus, unbridled, in refer- ence to the ability of this bacteriophage to attack all tested strains of the cholera organism. Common name : Cholera bacterio- phage C . Host : Vibrio comma Winslow et al., all strains. Geographical distribution: India. Induced disease : Sometimes death without lysis. When lysis occurs, it is rarely complete and is followed by sec- ondary resistant growth. Serological relationships : Antigenically distinct from cholera bacteriophage A. Immunological relationships : Second- ary growth resistant to this bacterio- phage, but susceptible to cholera bac- teriophages A and D. Other properties : Selectively inacti- vated by lipoid from smooth strain of host, but not by specific polysaccharide. Active in dilution of 1:10' or 1:10^". Multiplication rate, n X 10^ in 2 hours. Literature: Asheshov et al., Indian Jour. Med. Res., 20, 1933, 1127-1157; White, Jour. Path, and Bact., 43, 1936, 591-593. 44. Phagus lentus H. (loc. cit., 166). From Latin lentus, slow, in reference to the relatively slow and incomplete lysis induced by this bacteriophage. Common name : Cholera bacterio- phage D. Host : Vibrio comma Winslow et al . Geographical distribution : India. Induced disease : Incomplete lysis in about 5 hours, followed, in rough cultures, by slow development of resistant sec- ondary growth. Immunological relationships : Second- ary growth resistant to this bacterio- phage, but susceptible to cholera bac- teriophages A and C. Other properties : Not inactivated by specific polysaccharide effective against cholera bacteriophage A, nor by lipoid effective against cholera bacteriophage C. Multiplication rate, n X 10^ in 2 hours. Literature: Asheshov et al., Indian Jour. Med. Res., 20, 1933, 1127-1157; White, Jour. Path, and Bact., 43, 1936, 591-593. 45. Phagus diphtheriae H. {loc. cit., 167). From name of host. Common name : Corynebacierium diph- theriae bacteriophage. Host : Corynebacterium diphtheriae Lehmann and Neumann, many strains, especially 122 of 127 Australian type II gravis isolates; type I gravis isolates are Ij^sogenic (carriers) ; all intermediate iso- lates are susceptible. Insusceptible species : Corynebacterium diphtheriae, all tested mitis isolates, ex- cept 2 lysogenic. A strain of C. diph- theriae from Swan Hill, 200 miles north of Melbourne, was found to be resistant to this bacteriophage and to the small- 1144 MANUAL OF DETERMINATIVE BACTERIOLOGY plaque diphtheria bacteriophage, P. Ju- tilis. Geographical distribution : Australia. Induced disease : In Corynehacterium diphiheriae on agar, plaques 1 .0 to 1 .5 mm in diameter, with shelving edge. A few resistant bacterial colonies often appear in the central clear area. Literature: Keogh et al.. Jour. Path, and Bact., 46, 1938, 565-570; Smith and Jordan, Jour. Bact., SI, 1931, 75-88; Stone and Hobby, Jour. Bact., 27, 1934, 403-417. 46. Phagus futilis H. {loc. cit., 168). From Latin Jutilis, vain, in reference to regular appearance of resistant organisms in plaques on agar cultures lysed by this bacteriophage. Common name : Small-plaque diph- theria bacteriophage. Host : Corynehacterium diphtheriae Lehmann and Neumann, gravis type I isolates and all but 5 gravis type II iso- lates. Insusceptible species : All tested inter- mediate and mitis strains of C. diph- theriae. Geographical distribution : Australia. Induced disease : In Corynehacterium diphtheriae on agar, pin-point plaques or confluent plaques, with confluent growth of secondary, resistant organisms. Literature: Keogh et al.. Jour. Path, and Bact., ^6, 1938, 565-570. FAMILY CHLOEOGENACEAE 1145 SuBORDEK II. Phytophagineae suboirlo novus. Viruses infecting higher plants; vectors typically homopterous or hemipterous insects (leafhoppers, aphids, white flies, true bugs) or thysanopterous insects (thrips). From Greek phagein, to eat, and phyton, a plant. Key to the families of suborder Phytophagineae. 1. Inducing j-ellows-tj^pe diseases; vectors t^'pically cicadellid or fulgorid leaf- hoppers. Family I. Chlorogenaceae, p. 1145. 2. Inducing mosaic diseases ; vectors typically aphids. Family II. Marmoraceae, p. 1163. 3. Inducing ringspot diseases; vectors unknown. Family III. Annulaceae, p. 1212. 4. Inducing leaf -curl diseases; vectors typically white flies. Family IV. Rugaceae, p. 1218. 5. Inducing leaf-savoying diseases; vectors, true bugs. Family V. Savoiaceae, p. 1221. 6. Inducing spotted wilt; vectors, thrips. Family VI. Lethaceae, p. 1223. FAMILY I. CHLOROGENACEAE HOLMES EMEND. (Handb. Phytopath. Viruses, 1939, 1.) Viruses of the Yellows-Disease Group; pathogenic in flowering plants, causing dis- eases in which effects on chlorophyll are usually diffuse or stripe-like, no typical spot- ting or spotty mottling being involved. Vectors, so far as known, leafhoppers (CICA- DELLIDAE and F ULGORIDAE) . Key to the genera of family Chlorogenaceae. I. True Yellows Group. Viruses inducing diseases usually characterized by stimulation of normally dormant and adventitious buds to produce numer- ous slender shoots with long internodes and by chlorosis without spotting; invaded parts abnormallj' erect in habit. Vectors cicadellid leafhoppers so far as known . Genus I. Chlorogenns, p. 1146. II. Peach X-Disease Group. Viruses inducing diseases characterized by reset- ting of foliage and sometimes death of host. Genus II. Carpophthora, p. 1151. III. Phloem-Necrosis Group. Viruses inducing diseases characterized by progressive degeneration of the host plant or by wilting and sudden death ; sometimes by root discoloration. Vectors cicadellid leafhoppers so far as known. Genus III. Morsus, p. 1153. IV. Yellow-Dwarf Group. Viruses inducing diseases characterized by chlorotic effects somewhat resembling true mottling but often more diffuse. Vectors cicadellid (agallian) leafhoppers. Genus IV. Aureogenus, p. 1154. V. Fiji-Disease Group. Viruses inducing diseases characterized by marked vascular proliferation. The vector of one is known to be a leafhopper of the subfamily Delphacinae, family FULGORIDAE. Genus V. Galla, p. 1157. 1146 MANUAL OF DETERMINATIVE BACTERIOLOGY VI, Stripe-Disease Group. Viruses inducing diseases characterized by chlorotic striping; hosts grasses. Vectors, cicadellid and fulgorid leafhoppers. Genus VI. Fractilinea, p. 1159. Genus I. Chlorogenus Holmes. {Loc. cit., 1.) Viruses of the Typical Yellows Group, inducing diseases usually characterized by stimulation of normally dormant and adventitious buds to produce numerous slender shoots with long internodes, by chlorosis without spotting, or by both growth of nu- merous slender shoots and chlorosis. Invaded parts abnormally erect in habit. Af- fected flowers often virescent. Hosts, dicotyledonous plants. Vectors, so far as known, exclusively cicadellid leafhoppers. Generic name from Greek chloros, light green or yellow, and suffix, gen, signifying producing, from Greek genos, descent. The type species is Chlorogenus callistephi Holmes. Key to the species of genus Chlorogenus. I. Natural hosts many, in various families of plants. 1. Chlorogenus callistephi. 2. Chlorogenus australiensis. II. Known natural hosts relatively few. A. Natural hosts rosaceous. B. Natural hosts solanaceous. C. Natural host sandal. D. Natural host cranberry. E. Natural host locust. F. Natural host alfalfa. G. Natural host hop. 1. Chlorogenus callistephi Holmes. (Handb. Phytopath. Viruses, 1939, 2.) From New Latin Callistephiis, generic name of the China aster. Common names: Aster-yellows virus, lettuce white-heart virus, Erigeron-ycl- lows virus. Hosts : Callistephus chinensis Nees, the China aster, is the host that has been studied most. 170 or more species in 38 different families of dicotyledonous plants have been shown susceptible. Lettuce, endive, carrot, buckwheat, pars- nip, and New Zealand spinach are among the hosts of economic importance. 3. Chlorogenus persicae. 4. Chlorogenus solani. 5. Chlorogenus santali. 6. Chlorogenus vaccinii. 7. Chlorogenus robiniae. 8. Chlorogenus medicaginis. 9. Chlorogenus humuli. Insusceptible species : All tested spe- cies of the family Leguminosae and some species of all other tested families have appeared naturally immune. Geographical distribution: U. S., Can- ada, Bermuda, Japan, and Hungary. In California the celery-yellows strain of this virus replaces the type. Induced disease : In most host species the characteristics of disease are clearing of veins, followed by chlorosis of newly formed tissues, stimulation of normally dormant buds to growth, malformation, virescence of flowers, sterility, and up- right growth habit. Stimulation of nor FAMILY CHLOROGENACEAE 1147 mally dormant buds to adventitious growth and abnormally erect habit are the most constant features. Chlorosis is absent or inconspicuous in some hosts. Transmission : By leafhopper, Macro- steles divisus (Uhl.) (= Cicadiila sex- notata (Fall.), C. divisa (Uhl.)) {CICA- DELLIDAE). Incubation period, about 2 weeks . Some strains of this virus are transmitted also by the leafhoppers Thamnotettix montanus Van D. and T. geminatiis Van D. (CICADELLIDAE). By grafting. By dodder. Not through seeds of diseased plants. Not by me- chanical inoculation of plants, but virus has been passed from insect to insect mechanically in Macrosteles divisus; juice from viruliferous insects contains little virus just after inoculation, but the effec- tive concentration increases at least 100-fold between the 2nd and 12th day of a 17-day incubation period; it seems greatest before the insects begin to infect the aster plants on which they are main- tained. Thermal inactivation : In juice from viruliferous insects, at about 40° C in 10 minutes; at 25° C in 2 to 3 hours. In plant tissues, at 38° to 42° C, in 2 to 3 weeks; cured plants fully susceptible to reinfection. In insect vector, M. divisus, at 31° C in 12 days. Other properties : Virus in juices de- rived from insects is more stable at 0° C than at 25° C or when frozen ; at 0° C it withstands storage 24, not 48, hours in 0.85 per cent NaCl solution at pH 7.0 but most of the virus is inactivated in this time ; it withstands dilution 1:1000 in neutral 0.85 per cent NaCl solution; for brief (less than 5-minute) exposures, it re- mains viable over the range from pH 5 to 9. Literature: Black, Phytopath., SI, 1941, 120-135; 83, 1943, 2 (Abst.) ; John- son, ibid., 31, 1941, 649-656; Kunkel, Am. Jour. Bot., 13, 1926, 646-705; Contrib. Boyce Thompson Inst., 3, 1931, 85-123; 4, 1932, 405-414; Am. Jour. Bot., 2^, 1937, 316-327; 28, 1941, 761-769; Linn, Cornell Agr. Exp. Sta. (Ithaca), Bull. 742, 1940; Ogilvie, Bermuda Dept. Agr., Agr. Bull. 6, 1927, 7-8 ; Severin, Hilgardia, 3, 1929, 543-583; Phytopath., 20, 1930, 920-921; Hilgardia, 7, 1932, 163-179; 8, 1934, 305-325, 339-361; Phytopath., 30, 1940, 1049-1051 ; Hilgardia, 14, 1942, 411- 440; Severin and Haasis, Hilgardia, 8, 1934, 329-335. Strains : Two variant strains, one found in nature, the other derived experiment- ally, have been given varietal names to distinguish them from the type variety, vulgaris H. {loc. cit., 2) : la. Chlorogenus callistephi var. califor- nicus H. {loc. cit., 3). From California, name of state in which this strain was first recognized. Common name: Celery-yel- lows strain of aster-yellows virus. Dif- fering from the type variety by ability to infect celery {Apium gravcolens L. — UMBELLIFERAE) and zinnia {Zinnia elegans Jacq.— COMPOS IT AE) (Kun- kel, Contrib. Boyce Thompson Inst., 4, 1932, 405-414; Severin, Hilgardia, 3, 1929, 543-583; 8, 1934,305-325). lb. Chlorogenus callistephi var. ai- tenuatus H. {loc. cit., 4). From Latin attenuatus, weakened. Common name: Heat -attenuated strain of aster-yellows virus. Differing from the type variety by inducing less severe chlorosis and less uprightness of new growth in affected as- ter plants (Kunkel, Am. Jour. Bot., 2^, 1937,316-327). 2. Chlorogenus australiensis comb, nov. From Australia, name of continent. Synonym: Galla australiensis H. {loc. cit., 107). Common names : Tomato big-bud virus ; virescence virus; perhaps also stowboor virus, tobacco stolbur or montar virus, eggplant little-leaf virus. Hosts: SOLA N ACE AE— -Datura stra- vioniiim L., Jimson weed; Lycopersicon esculentum Mill., tomato ; Nicotiana taba- curn L., tobacco; Solaman melongena L., eggplant ; S. nigrum L., black nightshade. Recently a long list of species in this and 1148 MANUAL OF DETERMINATIVE BACTERIOLOGY other families have been reported as sus- ceptible to virescence virus, presumed to be an isolate of tomato big-bud virus. (Hill, Jour. Counc. Sci. Ind. Res., 16, 1943, 85-92). Geographical distribution : Australia, especially New South Wales; viruses causing somewhat similar diseases have been reported also from the Crimea and the northwestern United States. Induced disease: In tomato, flowers erect, virescent, calyx bladder-like, pol- len sterile; floral proliferation. Growth of axillary shoots stimulated. New leaves progressively smaller. Youngest leaves yellowish-green in color, especially at their margins ; usually purplish under- neath. Hypertrophy of inner phloem. No intracellular inclusions. Fruit red- dens imperfectly and becomes tough and woody. Roots appear normal. In So- lanum nigrum, axillary shoots numerous, leaves small, internal phloem adventi- tious. In tobacco, plants dwarfed ; leaves recurved, distorted, twisted, thickened, brittle, yellowish green, hanging down close to stem ; small leaves on shoots from axillary buds; proliferation and vires- cence of flowering parts ; chlorotic clear- ing of veins as early effect of disease ; upper surface of foliage appears glazed; some necrosis of veins, in old leaves, near tips and margins or on midrib; viable seed rarely produced; calyx bladder-like, floral axis may form short branches bear- ing small leaves ; disease sometimes called bunchy top. Transmission: By leafhopper, Tham- notettix argentata Evans {CICADELLI- DAE). Experimentally bj' budding and other methods of grafting. Not by inocu- lation of expressed juice. Literature : Cobb, Agr. Gaz. New South Wales, 13, 1902, 410-414; Dana, Phyto- path., 30, 1940, 866-869; Hill, Jour. Aus- tral. Inst. Agr. Sci., 6, 1940, 199-200; Jour. Council Sci. Ind. Res., 10, 1937, 309-312; 16, 1943, 85-92; Michailowa, Phytopath., 25, 1935, 539-558; Rischkov et al., Ztschr. Pflanzenkr., 43, 1933, 496- 498; Samuel et al., Phytopath., 23, 1933, 641-653. 3. Chlorogenus persicae H. (loc. cit., 5). From New Latin Persica, former generic name of peach. Common names : Peach-yellows virus, little-peach virus. Hosts : ROSACE AE — Prunus persica (L.) Batsch, peach; P. salicina Lindl., Japanese plum; and all other tested spe- cies of the genus Prunus. Geographical distribution : Eastern United States and Canada, south to North Carolina. First occurred near Philadelphia in this country. Origin perhaps oriental ; introduction in oriental plums suspected. Not in Europe. Induced disease : In peach, clearing of veins, production of thin erect shoots bearing small chlorotic leaves, followed by death in a year or two . In early stages of the disease there is premature ripening of fruit. In Japanese plum, systemic in- fection but no obvious symptoms. Transmission : By the leafhopper, Ma- cropsis trimaculala (Fitch) {CICADEL- LIDAE). By budding; virus spreads down stem from point of bud insertion faster than up. Not by inoculation of expressed juice, despite numerous at- tempts. Not by pollen of diseased trees. Immunological relationships : Presence of peach-yellows virus immunizes tree against little-peach virus, formerly con- sidered an independent entity. Thermal inactivation : In peach tissues, at 34° to 35° C in 4 to 5 days ; at 44° C in 30 minutes ; at 47° C in 10 minutes ; at 50° C in 3 to 4 minutes ; at 56° C in 15 seconds. Other properties : Trees and bud sticks may be treated safely with heat sufficient to kill the virus. Cured trees are sus- ceptible to reinfection. Literature : Blake, N. J. Agr. Exp. Sta., Bull. 226, 1910; Kunkel, Contrib. Boyce Thompson Inst., 5, 1933, 19-28; Phyto- path., 26, 1936, 201-219, 809-830; 28, 1938, 491-497; Manns, Trans. Peninsula Hort. Soc, 23, 1933, 17-19; Manns and Manns, FAMILY CHLOROGENACEAE 1149 ibid., 24, 1934, 72-76; McCubbin, Penn- sjdvania Dept. Agr., Gen. Bull. 382, 1924. Strains : Numerous strains of peach- yellows virus probably exist in nature. One of these has been given a varietal name, distinguishing it from the tj^pe variety, vulgaris H. (loc. cit., 5) : 3a. Chlorogenus ■persicae var. microper- sica H. {loc. cit., 6). From Greek micros, small, and New Latin Persica, former generic name of peach. Common name : Little-peach strain of peach-yellows virus. Differing from the type variety by tend- ency to cause a mild type of disease, characterized by distortion of young leaves, production of many short branches on main trunk, later yellowing of mature leaves, twiggy growth, shoots slightly less erect than in typical peach yellows. (Kunkel, Phytopath., 26, 1936, 201-219; 26, 1936, 809-830; 28, 1938, 491-497; Manns, Trans. Peninsula Hort. Soc, 23, 1933, 17-19; 24, 1934, 72-76.) 4. Chlorogenus solani H. {loc. cit., 7). From New Latin Solanum, generic desig- nation of potato. Synonym: Chloroph- thora solani McKinney, Jour. Washington Acad.Sci., 34,1944,151. Common names : Potato witches'-broom virus, potato wilding or semi -wilding virus. Hosts: SOL AN AC EAE— Solanum tu- berosum L., potato. Experimentally, also SOLA NACEAE — Lycopersicon esculen- tum Mill., tomato ; Nicotiana tabacwn L., tobacco; A'^. glutinosa L.; N. rusiica L. APOCYNACEAE—Vinca rosea L., periwinkle. CHENOPODIACEAE— Beta vulgaris L., sugar beet. Geographical distribution : United States (Montana, Washington), Russia. Induced disease: In potato, increas- ingly pronounced flavescence in new leaflets on one or more stems, production of new dwarfed leaflets with marginal flavescence on stems with unusually long internodes and enlarged nodes, growth of spindling axillary and basal branches. profuse blooming and fruiting, lack of dormancy in tuber buds, formation of many small underground tubers as well as some aerial tubers ; plants grown from diseased tubers form thread-like stems and small simple leaves; infected plants survive several seasons, with progressive degeneration. In tomato, experimen- tally, extreme leaf dwarfing, marginal flavescence of leaves and abnormally numerous axillary branches; stems be- come hollow and die ; plants do not survive long after infection. In tobacco, experi- mentally, slender axillary branches with dwarfed leaves, flowers on spindling pedi- cels, numerous, small ; later leaves flaves- cent or marginally flavescent. Transmission : By tuber-core grafts with pre patent period of 29 to 114 days. By stem grafts. By dodder, Cuscuta campestris Yuncker {CONVOLVULA- CEAE). Not by inoculation of ex- pressed juice. Not by Macrosteles divi- sus (Uhl.) {CICADELLIDAE). No insect vector is known. Not through seeds of diseased tomatoes. Thermal inactivation : at 42° C in 13 days, in tissues of Vinca rosea; at 36° C in 6 days in small potato tubers. Literature : Hungerford and Dana, Phytopath., I4, 1924, 372-383; Kunkel, in Virus Diseases, Cornell Univ. Press, Ithaca, N. Y., 1943, 63-82; Proc. Am. Philosoph. Soc, 86, 1943, 470-475; Mc- Larty, Scient. Agr., g, 1926, 395; Whipple, Montana Agr. Exp. Sta., Bull. 130, 1919; Young, Science, 66, 1927, 304-306; Am. Jour. Bot., 16, 1929, 277-279; Young and Morris, Jour. Agr. Res., 36, 1928, 835-854. 5. Chlorogenus santali H. {loc. cit., 8). From New Latin Santalum, generic desig- nation of sandal. Common names : Sandal spike -disease virus, sandal spike-rosette virus. Hosts: S AN TAL ACE AE— Santalum album, L., sandal. Spike-like diseases have been found also in RHAMNEAE— Zizyphus oenopUa Mill., SAPINDA- CEAE — Dodonaea viscosa Jacq., VER- 1150 MANUAL OF DETERMINATIVE BACTERIOLOGY BENACEAE — Stachytarpheta indica Vahl, and APOCYN ACEAE—Vinca ro- sea L. Geographical distribution : South India. Induced disease : In sandal, abnormally profuse blooming at first, suppression of blooming later; reduction in leaf size and internode length; death ensues in the third year or earlier. In all but the youngest leaves of affected branches, vacuolate intracellular bodies with defi- nite peripheral membrane, 4 to 9 microns in maximum diameter, are found. Transmission: By twig grafts, inserted buds, and patch grafts, with success de- creasing in the order named. Prepatent period 3 to 4 months. Best results in May and June ; poorest in October. Per- haps through seeds, but not through pollen of diseased plants. Insect trans- mission claimed, but species not identi- fied. Reported transmission by Moonia alhimaculata (CICADELLIDAE) re- quires further confirmation. Not by inoculation of expressed juice. Not by root grafts. Literature : Coleman, Mysore Dept. Agr., Mycol. Ser., Bull. 3, 1917; Indian Forester, 49, 1923, 6-9; Dover, ibid., 60, 1934, 505-506; Narasimhan, Phytopath., 23, 1933, 191-202 ; Rangaswami and Sreen- vasaya. Current Science, 4, 1935, 17-19; Sreenvasaya, Nature, 126, 1930, 957; Venkata Rao and Gopala Iyengar, Mysore Sandal Spike Invest. Comm., Bull. 4, 1934, 1-12; Indian Forester, 60, 1934, 689-701. 6. Chlorogenus vaccinii H. {loc. cit., 10) . From New Latin Vacciniuvi , generic designation of cranberry. Common names : Cranberry false-blos- som virus, Wisconsin false-blossom virus. Hosts : ERICACEAE — Vaccinium nia- crocarpon Ait., cranberry; V. oxycoccus L. Experimentally, also APOCYNA- CEAE — Vinca rosea L., periwinkle. COMPOSIT AE—Calend\ila officinalis L., calendula. SOLAN ACE AE—Lyco- persicon esculentuvi Mill., tomato; Nico- tiana glutinosa L.; N. tabacum L., to- bacco; Solanum tuberosum L., potato. Geographical distribution : Eastern United States and Canada. It is be- lieved that the virus does not spread in bogs with alkaline (pH 7.4 to 8.8) flooding water in Wisconsin though it spreads rapidly in the more productive bogs with nearly neutral (pH 6.0 to 7.0) flooding water. Induced disease : In cranberry, flowers erect, instead of pendent as in healthy plants; calyx lobes enlarged, petals short, streaked with red and green, stamens and pistils abnormal. Flowers may be re- placed by leaves or short branches. Dor- mancy of axillary buds is broken, pro- ducing numerous erect shoots, forming a witches' broom. Diseased fruits small, irregular in shape, erect. Transmission : By leafhopper, Ophiola striatula (Fall.) ( = Euscelis striatulus (Fall.)) {CICADELLIDAE). Not by inoculation of expressed juice. By dod- der, Cuscuta campestris Yuncker (CON- VOLVULACEAE). Thermal inactivation : At 40° C in 2 weeks in tissues of periwinkle. Literature : Dobroscky, Contrib. Boyce Thompson Inst., 3, 1931, 59-83; Fracker, Phytopath., 10, 1920, 173-175; Kunkel, Science, 95, 1942, 252; Torreya, 43, 1943, 87-95; Shear, U. S. Dept. Agr., Bull. 444, 1916; Stevens, Phytopath., 15, 1925, 85- 91; 34, 1944, 140-142; U. S. Dept. Agr., Circular 147, 1931 ; Stevens and Sawyer, Phytopath., 16, 1926, 223-227 ; Wilcox and Beckwith, Jour. Agr. Res., 47, 1933, 583-590. 7. Chlorogenus robiniae H. (loc. cit., 13). From New Latin Robinia, generic designation of locust. Synonym: Poly- cladus robiniae McKinney, Jour. Wash- ington Acad. Sci., 34, 1944, 151. Common names: Locust witches '- broom virus; locust brooming-disease virus. ■ Hosts: LEGUM I NOSAE— Robinia pseudoacacia L., black locust. Geographical distribution : United FAMILY CHLOROGENACEAE 1151 States (southern Pennsylvania to north- eastern Georgia, west to southwestern Ohio and Tennessee). Induced disease : In black locust, clear- ing of veins, followed by progressive re- duction in size of newly formed leaves; growth of spindly shoots to form witches' brooms. Roots more brittle, shorter, and darker than normal, with excessive branching of rootlets, giving the appear- ance of root brooms. Transmission : By grafting and bud- ding. Not by inoculation of expressed juice. No insect vector is known. Literature :Grant etal., Jour. Forestry, 40, 1942, 253-260; Hartley and Haasis, Phytopath., 19, 1929, 163-166; Jackson and Hartley, Phytopath., 23,193S, 83-9 0; Waters, Plant World, 1, 1898, 83-84. 8. Chlorogenus medicaginis H. {loc. cil., 14). From New Latin Medicago, generic designation of alfalfa (lucerne). Common names : Alfalfa witches'- broom virus, lucerne witches'-broom virus, spindle-shoot virus, mistletoe virus, Kurrajong virus, bunchy-top virus. Hosts : LEG UM I NOS A E— Medicago sa(it;a L., alfalfa (lucerne). Geographical distribution : Australia, especially New South Wales; perhaps United States. Induced disease : In alfalfa, plant dwarfed; leaves small, rounded, chlorotic at edge, puckered, distorted ; stems short, spindly, numerous. Flowers usually not formed, but sometimes pale and small, sometimes replaced by leafy structures. Seed rarely produced. Transmission: By grafting. Not by inoculationof expressed juice. No insect vector is known. Literature : Edwards, Jour. Australian Inst. Agr. Sci.,1, 1935, 31-32; New South Wales Dept. Agr., Science Bull. 52, 1936; Agr. Gaz. New South Wales, 47, 1936, 424-426; Richards, U. S. Dept. Agr., Plant Disease Reporter, Supplement, 71, 1929, 309-310. 9. Chlorogenus humuli H. {loc. cit., 15). From New Latin Humulus, generic designation of the hop. Common names : Hop-nettlehead virus, silly-hill disease virus, virus of infectious- sterility of the hop. Hosts : MO RACE AE— Humulus lupu- lus L., European hop. Geographical distribution : England, Czechoslovakia, Germany, Poland. Induced disease : In hop, stems numer- ous, spindly, short, plants weak. Leaves curled upward at margin; cone produc- tion greatly reduced. Transmission: By grafting. Not by in- oculation of expressed juice. Not through soil. No insect vector is known. Literature : Blattny and Vukolov, Rec. Inst. Rech. Agron. Rep. Tchecosl., 137, 1935, 3-18; Goodwin and Salmon, Jour. Inst. Brew., 33, 1936, 209-210; Salmon, ibid., 32, 1935, 235-237; 33, 1936, 184-186; Salmon and Ware, Jour. South-Eastern Agr. College, Wye, Kent. 37, 1936, 21-25. Genus II. Carpophthora Mc Kinney emend. (Jour. Washington Acad. Sci., 34, 1944, 152.) Peach X-Disease Group; viruses inducing diseases characterized in general by rosetting of foliage and sometimes death of host. Generic name from Greek, meaning fruit and ruin or destruction. The type species is Carpophthora lacerans McKinney. Key to the species of genus Carpophthora. I. Inducing chlorosis, reddening, and tattering of foliage, with rosette formation late in the disease in some hosts. 1. Carpophthora lacerans. 1152 MANUAL OF DETERMINATIVE BACTERIOLOGY II. Inducing rosette formation characteristically, but not tattering of affected foliage. 2. Carpophthora rosettae. 1. Carpophthora lacerans (Holmes) McKinney. {Marmor lacerans Holmes, Handb. Phytopath. Viruses, 1939, 82; McKinney, Jour. Washington Acad. Sci., 34, 1944, 152.) From Latin lacerare, to lacerate, in reference to characteristic foliage injury. Common name : Peach X-disease virus ; virus of peach yellow-red virosis. Hosts: ROSACE AE — Primus persica (L.) Batsch, peach; P. virginiana L., chokecherry. Geographical distribution : United States, Canada. Induced disease : In peach, foliage nor- mal each spring but yellowish areas ap- pear in June at base of leaves ; affected trees appear lighter green than neighbor- ing healthy trees; discolored spots occur at random on the leaf blade, becoming red and yellow with remainder of leaf becom- ing chlorotic ; the discolored areas usually fall out, leaving the foliage tattered ; sub- sequently, affected leaves drop except at tips of branches; young trees may die, older ones survive indefinitely. Fruit either shrivels and falls or ripens prema- turely, with bitter flavor and no viable seed. In chokecherry, conspicuous pre- mature reddening of foliage, development of fruits with dead embryos in the pits ; in the second and third seasons after infection, duller colors of foliage, rosettes of small leaves on terminals; death fol- lows the advanced stage of disease. Transmission : By budding. Not bj' inoculation of expressed juice. No insect vector has been reported. Literature: Berkeley, Div. of Botany and Plant Path., Science Service, Do- minion Dept. Agr., Ottawa, Canada, Pub- lication 678, 1941 ; Boyd, U.S. Dept. Agr., Plant Dis. Rep., 22, 1938, 334; Hilde- brand, Contrib. Boyce Thompson Inst., 11, 1941, 485-496; Hildebrand and Pal- miter, U. S. Dept. Agr., Plant Dis. Rep., 22, 1938, 394-396; Hildebrand et al.. Handbook of virus diseases of stone fruits in North America, Michigan Agr. Exp. Sta., Misc. Publ., 1942, 21-24; Stoddard, Connecticut Agr. Exp. Sta., Circ. 122, 1938, 54-60; Proc. Connecticut Pomol. Soc, 48, 1938, 29-32. 2. Carpophthora rosettae (Holmes) comb. nov. {Cfdorogeniis rosettae H., nomen nudum, Phytopath. 29, 1939, 434; Nanus rosettae H., Handb. Phytopath. Viruses, 1939, 125.) From rosette, com- mon name of induced disease, from French, diminutive of rose, a rose. Common name: Peach-rosette virus. Hosts: ROS ACE AE—Prunus persica (L.) Batsch, peach; P. communis Fritsch, almond; P. domestica L., plum. Experi- mentally, SiXso—APOCYNACEAE— Vinca rosea L., periwinkle. ROSA- CEAE — P. am errcana Marsh., wild plum ; P. armeniaca L., apricot; P. cerasus L., cherry; P. pumila L., sand cherry. SOLA NACEAE — Lycopersicon esculen- tum Mill., tomato ; Nicotiana glvtinosa L. Geographical distribution : United States (Georgia, Alabama, South Caro- lina, Tennessee, West Virginia, Missouri, Oklahoma). Induced disease : In peach, sudden wilting and death, or growth of abnor- mally short stems bearing dwarfed leaves with clearing and thickening of veins, followed by death in a few months. Transmission : By budding. By dod- der, Cuscuta campestris Yuncker. Not by inoculation of expressed juice. Not through soil. No insect vector is known. Immunological relationships : No pro- tection is afforded by previous infection of peach trees with Chlorogenus persicae, peach-yellows virus. Thermal inactivation : At 50° C in 10 minutes (in tissues of peach). Rosetted trees are abnormally susceptible to heat FAMILY CHLOROGENACEAE 1153 injury and heat treatments cure peach- rosette disease only in recently infected trees. Literature: Kunkel, Phytopath., 26, 1936, 201-219, 809-830; in Virus Diseases, Cornell Univ. Press, Ithaca, N. Y., 1943, 63-82; McClintock, Jour. Agr. Res., U, 1923, 307-316; Phytopath., 21, 1931, 373- 386; Smith, U. S. Dept. Agr., Div. Veg. Path., Bull. 1, 1891. Genus III. Morsus gen. nov. Alfalfa-Dwarf Group; viruses inducing diseases characterized in general by sud- den wilting and death or by gradual decline of vigor with foliage of darker green color than normal. Vectors, like those of the typical yellows subgroup, cicadellid leafhoppers so far as known. Generic name from Latin morsus, sting or vexation. The type species is Morsus suffodiens spec. nov. Key to the species of genus Morsus. I. Affecting alfalfa and grape. 1. Morsus suffodiens. II. Affecting tobacco. 2. Morsus reprimens. III. Affecting elm. 3. Morsus ulmi. 1. Morsus suffodiens spec. nov. From Latin suffodere, to sap or undermine, in reference to process leading to sudden collapse of long infected, but sometimes not obviously injured, grape vines as well as to progressive decline in size of in- fected alfalfa plants, the foliage of which may remain green to the last. Common names: Alfalfa-dwarf virus, lucerne dwarf -disease virus, virus of Pierce's disease of the grape, virus of Anaheim disease. Hosts : LEG UM INOSAE—Medicago sativa L., alfalfa (lucerne). VITA- CEAE — Vitis vinifera L., grape. Geographical distribution: United States. Induced disease : In alfalfa, leaves small but green, plant progressively smaller, wood of I'oots discolored yellow, transpiration decreased; wilting may oc- cur; starch of root diminished; plant eventually succumbs, thinning stand prematurely. In grape, dark green color of leaves retained along veins, not be- tween them, or no abnormality in appear- ance of foliage ; wilting and sudden death of plant in summer of second year. In late summer of first year, there may be dying leaf margins and dying back of cane tips. Transmission : By budding and root grafting. By leafhoppers, Draeculace- phala minerva Ball, Carneocephala Julgida Nott., C. trigultata Nott., Helochara delta Oman, Neokolla circellata (Baker), N. gothica (Sign.), N. confluens (Uhler), N. heiroglyphica (Say), and Cuerna oc- cidentalis Oman and Beamer (CICA- DELLIDAE) ; these vectors all belong to the subfamily Amhlycephalinae; all tested species of this, but none of any other, subfamily have proved capable of transmitting this virus. Not by in- oculation of expressed j uice . Not through soil. Literature: Frazier, Phytopath., 3^, 1944, 1000-1001; Hewitt, Phytopath., 29, 1939, 10; SI, 1941, 862; Blue Anchor, 18, 1941, 16-21, 36; Hewitt et al., Phytopath., 32, 1942, 8; Houston et al., Phytopath., 32, 1942, 10; Milbrath, Calif. Dept. Agr., 20th Ann. Rept., Bull. 28, 1940, 571; Pierce, U. S. Dept. of Agr., Div. of Veg. Path., Bull. 2, 1892, 1-222; Weimer, Phytopath., 21, 1931, 71-75; 27, 1937, 697-702; Jour. Agr. Res., 47, 1933, 351- 368; 53, 1936, 333-347; 55, 1937, 87-104. 1154 MANUAL OF DETERMINATIVE BACTERIOLOGY 2. Morsus reprimens .spec. nov. From Latin reprimere, to restrain, check, or curb, in reference to the inhibiting effect on growth of the host plant, tobacco. Common name : Tobacco yellow-dwarf virus. Hosts: SOLAN ACE AE—Nicoiiana tabacum L., tobacco; A'", rustica L., In- dian tobacco; N. trigonophylla Dun. Experimentally, also N. glauca Grah. (symptomless) and N. glutinosa L. Geographical distribution : Australia (Victoria, New South Wales, South Aus- tralia, and southern Queensland). Induced disease : In tobacco, internodes of stem shortened, leaves small; down- ward bending of tips and rolling under of margins of young apical leaves; young leaves darker than normal at first, bunched, later appear ribbed; leaves be- come yellow-green, pale first between veins; old leaves rugose, thickened, later savoyed. Root system small, roots slightly brown externally and in the re- gion of the phloem. Affected plants may survive the winter and show diseased new growth in the spring. Transmission: By grafting and bud- ding. By nymphs and adults of the leaf- hopper Thamnoiettix argentata (Evans) (CICADELLIDAE). Literature : Dickson, Australia, Coun- cil Sci. Indust. Res., Pamphlet 14, 1929, 22; Hill, Australia, Journal of the Coun- cil Sci. Indust. Res., 10, 1937, 228-230; 14, 1941, 181-186; 16, 1942, 13-25. 3. Morsus ulmi spec. nov. From Latin ulmus , ehn. Common name : Elm phloem-necrosis virus. Host : U RT IC ACE AE— Ulmus ameri- cana L., American elm. Geographical distribution : United States (Ohio, Indiana, Illinois, Missouri, Tennessee, Kentucky, and West Vir- ginia). Induced disease: In elm, gradual de- cline over a period of 12 to 18 months before death or sudden wilt, drying of leaves, and death within 3 to 4 weeks. All ages susceptible, from seedling to large tree. Transmission : By patch grafting. Not by inoculation of expressed juice. Literature : Leach and Valleau, U. S. Dept. Agr., Plant Dis. Rept., 23, 1939, 300-301; Swingle, Phytopath., 30, 1940, 23. Genus IV. Aureogenus Black. (Proc. Am. Philos. Soc, 88, 1944, 141.) Viruses of the Yellow-Dwarf Group, inducing diseases characterized by yellowing without typical mosaic-type mottling. Vectors agallian leafhoppers (CICADELLI- DAE). Generic name from Latin aureus, yellow or golden, and genus, group. The type species is Aureogenus vastans (Holmes) Black. Key to the species of genus Aureogenus. I. Mechanically transmissible in some hosts by rubbing methods'of inoculation. Not producing enlarged veins or club-leaf in clover. 1. Aureogenus vastans. II. Not known to be transmissible by rubbing methods of inoculation. A. Producing enlarged veins in clover. 2. Aureogenus magnivena. B. Producing club-leaf in clover. 3. Aureogenus clavifolium. FAMILY CHLOROGEXACEAE 1155 1. Aureogenus vastans (Holmes) Black. {Mannor vastans Holmes, Handb. Phytopath. Viruses, 1939, 94; Black, Proc. Am. Philos. Soc, 88, 1944, 141.) From Latin vastare, to devastate. Common name : Potato yellow-dwarf virus. Hosts: SOLAN ACE AE—Solanum tu- berosum L., potato. COMPOSITAE— Chrysanthemum leucanthemum L., var. ■pinnatifidum Lecoq and Lamotte, daisy; Rudheckia hirta L., black-eyed Susan. CRUCIFERAE—Barbarea vulgaris R. Br., common winter cress. LEGU- MINOSAE — Trifolium pratense L., red clover. Experimentally to numerous species in these and other families. Geographical distribution: Northeast- ern United States and southeastern Canada. Induced disease : In potato, yellowing of leaves, necrosis of stem, dwarfing of plant; the stem, if split, shows rusty specks especially at nodes and apex ; the apex dies early; tubers are few, small, close to the stem, often cracked, with flesh discolored by scattered brown specks; seed tubers tend to remain un- rotted in the ground, becoming hard and glassy ; some of them do not germinate in warm soil, others produce shoots that die before reaching the surface, giving poor stands. In Chrysanthemum leucanthe- mum var. pinnatifidum, at first, clearing of veins; later, young leaves distorted, thick, stiff, small; petioles short, leaves erect, forming a rosette at the crown of the plant ; with passing of the early phases of the disease, foliage tends to appear nearly normal, but remains darker green and more erect than that of healthy plants; virus is recoverable both during and after the period of obvious disease and infected plants may constitute an important reservoir. In Trifolium in- carnatum L., crimson clover, experiment- ally, clearing of veins and yellowing of younger leaves (in summer the yellowing is usually replaced in part by an inter- veinal reddish-brown color on both leaf surfaces extending from the margins in- wards); dwarfing of entire plant; death or a chronic disease characterized by milder manifestations without, however, vein enlargement or cupping of leaves. In Nicotiana rustica L., experimentally, yellowish primary lesions followed by clearing of veins and systemic chlorosis; the primary lesions facilitate quantitative estimation of concentrations of this virus. Transmission : By inoculation of ex- pressed juice, in the presence of finely powdered carborundum, to Nicotiana rustica; mechanical transmission very difficult in other hosts tested. By graft- ing. By clover leafhopper, Aceratagallia sanguinolenta (Provancher) ; experiment- ally, by other closely related leafhoppers, Aceratagallia lyrata (Baker), A. obscura Oman, and A. curvata Oman; not (for the type variety of the virus) by Agallia con- stricta Van Duzee ; very rarely by Agallia quadripunctata (Provancher) and Agal- liopsis novella (Say) (CICADELLI- DAE). The vector Aceratagallia san- guinolenta remains infective as an over- wintering adult ; incubation period not less than 6 days, commonly much longer; virus does not pass to progeny of virulif- erous leafhoppers through eggs or sperm; this leafhopper varies genetically in ability to transmit. Immunological relationships : No pro- tection is afforded against necrotic effects of a testing strain of this virus (var. lethale Black) by prior inoculation of Nicotiana rustica with isolates of Marmor medicaginis, M. cucumeris, M. upsilon, Annulus tabaci, A. orae, or A. dubius, but the varieties vulgare Black and agal- liae Black protect ; these specifically pro- tecting strains give no similar protection against formation of necrotic lesions by subsequently applied isolates of Marmor tabaci, M. lethale, Annulus tabaci, or A. orae. Thermal inactivation : At 50 to 52° C in 10 minutes. 1156 MANUAL OF DETERMINATIVE BACTERIOLOGY Filterability : Passes Berkefeld W filter. Other properties : Virus viable at 23 to 27° C less than 13 hours after extraction of juice from diseased plant ; not infective after drying in leaf tissues. Literature : Barrus and Chupp, Phyto- path., 12, 1922, 123-132; Black, Am. Po- tato Jour., 11, 1934, 148-152; Cornell Univ. Agr. E.xp. Sta., Mem. 209, 1937, 1-23; Phytopath., £8, 1938, 863-874; Am. Jour. Bot., 27, 1940, 386-392; Am. Potato Jour., 18, 1941, 231-233; Phytopath., 33, 1943, 363-371; Genetics, 28, 1943, 200- 209; Proc. Am. Philos. Soc, 88, 1944, 132-144; Hansing, Cornell Univ. Agr. Exp. Sta., Bull. 792, 1943; Price and Black, Am. Jour. Bot., 28, 1941, 594-595; Taylor, Am. Potato Jour., 15, 1938, 37-40; Walker and Larson, Jour. Agr. Res., 59, 1939, 259-280; Wat kins. Jour. Econ. Ent., 32, 1939, 561-564 ; Cornell Univ. Agr. Exp. Sta., Bull. 758, 1941, 1-24; Younkin, Am. Potato Jour., 19, 1942, 6-11. Strains: Beside the type variety, Aureogerms vastans var. vulyare Black (Am. Jour. Bot., 27, 1940, 391), on which the species is based, two distinctive strains have been given varietal names : la. Avreogenus vastans var. agalliae Black. (Am. Potato Jour., 18, 1941, 233.) From New Latin Agallia, generic name of vector of this strain. Common name : New Jersey strain of potato yellow-dwarf virus. Differing from the type especially in its distinctive vector, the leafhopper, Agallia consiricta Van Duzee, which is incapable of transmitting the type strain, and in not being transmitted by Acerata- gallia sangidnolenta (Provancher), com- mon vector of the type variety. Experi- mentally, transmitted also by Agallia quadripunctata (Provancher) ; perhaps rarely by Agalliopsis novella (Say); Differing but little from the type in ef- fects on potato (var. Green Mountain) and Nicotiana rustica but more definitely in effects on crimson clover, in affected plants of which a rusty-brown necrosis along the veins, not induced by the type strain, is always present in some degree. lb. Aureogenus vastans var. lethale Black. (Am. Jour. Bot., 27, 1940, 391.) From Latin lethalis, causing death. Common name : Strain B5 of potato yel- low-dwarf virus . Differing from the type variety especially in a tendency to induce in Nicotiana rustica, experimentally, brown primary lesions with necrotic gray centers, systemic yellowing, extensive necrosis of veins, collapse of large areas of leaf, and sometimes death of the host ; not known to occur in nature as a separate strain, but readily isolated as a variant from strains collected in nature. 2. Aureogenus magnivena Black. (Proc. Am. Philos. Soc, 88, 1944, 144.) From Latin magnus, large, and vena, vein. Common name : Clover big-vein virus. Host: Experimentally, LEGUMINO- SAE — Trifolium incarnatum L., crimson clover. Insusceptible species: SOLAN A- CEAE — Nicotiana rustica L., Indian to- bacco; Solanum tuberosum L., potato. Geographical distribution : United States (presumably, Washington, D. C). Induced disease : In crimson clover, experimentally, unevenly thickened veins which are depressed below the up- per surface of the leaf; these enlarged veins, best observed from below, some- times bear enations that arise from their lower surfaces, leaves often curl upward and inward marginally; in summer, yel- lowing of leaves progresses from margins inward, the yellow color being later re- placed in part by red or purple red ; petioles undulating; plants dwarfed; internodes shortened; no clearing of veins; no rusty-brown necrosis. Transmission: Not by inoculation of expressed juice. By leafhoppers, Agal- liopsis novella (Say), Agallia constricta Van Duzee, A. quadripunctata (Provan- cher) ; not by Aceratagallia sangnino- lenia (Provancher) (CICADELLIDAE). FAMILY CHLOROGENACEAE 1157 3. Aureogenus clavifolium Black. (Proc. Am. Philos. Soc, 88, 1944, 141.) From Latin clava, club, a.nd folium, leaf. Common name : Clover club-leaf virus. Host: Experimentally, LEGUMIXO- SAE — Trifolium incarnatum L., crimson clover. Insusceptible species: SOLAX A- CEAE — Nicotiana rustica L., Indian to- bacco ; Solanum tuberosum L., potato. Geographical distribution : United States (Princeton, X. J.). Induced disease : In crimson clover, ex- ])erimentalh-, j-oungest leaves lighter green than normal, slow to unfold; leaf margins jellowed or colored red or purple red; affected leaves narrow, smooth or savoyed; plant dwarfed, new shoots from leaf axils slightly stimulated ; new growth of spindly stems and small leaves; no rusty-brown necrosis of veins, no obvious enlargement of veins, and no obvious clearing of veins at the onset of disease. Transmission : Xot by inoculation of expressed juice. By leafhopper, Agal- liopsis novella (Say) (CICADELLI- DAE) ; not by leafhoppers, Aceratagallia sanguinolenta (Provancher), Agallia con- stricia Van Duzee, nor ^4. quadripunctata (Provancher) (CICADELLIDAE). Genus V. Galla Holmes. (Loc. cit., 106) Viruses of the Fiji-Disease Group, inducing diseases characterized by vascular proliferation. Generic name from Latin galla, a gall nut. The type species is Galla fijiensis Holmes. Key to the species of genus Galla. I. Infecting sugar cane. A. Inducing formation of conspicuous galls. 1. Galla fijiensis. B. Xot inducing formation of conspicuous galls. 2. Galla queenslandiensis . II. Infecting anemone. 3. Galla anemones. III. Infecting peach. 4. Galla verrucae. IV. Infecting corn. 5. Galla zeae. 1. Galla fijiensis Holmes. (Handb. Phytopath. Viruses, 1939, 106.) From name of Fiji Islands. Common name: Fiji-disease virus. Host : GRAM I XEAE—Saccharum offi- cinarum L., sugar cane. Geographical distribution : Fiji Islands, X'ew South Wales, Java, Philippine Is- lands, Xew Guinea and Xew Caledonia. Induced disease: In sugar cane, galls on vascular bundles, formed by prolifera- tion of phloem and nearby cells. Af- fected cells show characteristic spherical or oval inclusion bodies. Developing leaves shortened, crumpled, abnormally dark green. Infected stools of cane be- come bush}^ Roots small, bunchy. Transmission: By leafhoppers, Perkin- siella saccharicida Kirk, (in Queensland) and P. vastatrix Breddin (in Philip- pine Islands) {FULGORIDAE, sub- family Delphacinae) . Xot by grafting. Xot by inoculation of expressed juice. Xot through eggs of P. vastatrix. Cut- tings taken from affected canes produce some healthy and some diseased plants, because virus does not become uniformly distributed throughout the host tissues. Literature: Kunkel, Bull. Exp. Sta., Hawaiian Sugar Planters' Assoc, Bot. 1158 MANUAL OF DETERMINATIVE BACTERIOLOGY Ser., 3, 1924, 99-107; Lyon, ibid., 3, 1921, 1-43; Hawaiian Planters' Rec, 12, 1915, 200; Mungomery and Bell, Queensland, Bur. Sugar Exp. Sta., Div. Path., Bull. 4, 1933; Ocfemia, Am. Jour. Bot., 21, 1934, 113-120 ; Ocfemia and Celino, Phyto- path., ^5, 1939, 512-517 ; Reinking, Phyto- path.,ii, 1921, 334-337. 2. Galla queenslandiensis H. {loc. cit., 109). From Queensland, where the in- duced disease was first studied. Common name : Sugar-cane dwarf -dis- ease virus. Host : GRAM INEAE—Saccharum offi- cinarum L., sugar cane. Geographical distribution : Queensland. Induced disease: In sugar cane, young leaves marked with scattered chlorotic streaks, leaves stiff and erect, spindle twisted, abnormally short and pale. As leaves mature, streaks disappear, leaves become darker than normal green. In recently infected plants, vascular bun- dles are enlarged, irregular in shape, fused and characterized by abnormal prolifera- tion of thin-walled lignified cells. Literature : Bell, Queensland, Bur. Sugar Exp. Sta., Div. Path., Bull. 3, 1932. 3. Galla anemones H. {loc. cit., 108). From Latin anemone, anemone or wind- flower. Common name : Anemone -alloiophylly virus. Hosts: RANUNCULACEAE— Ane- mone nemorosa L., vernal windflower; A. ranunculoides L. ; .A. trifolia L. Geographical distribution: Germany. Induced disease : Leaves thickened and distorted, petioles thickened. Flowers distorted or not formed. Vascular bun- dles larger and more numerous than in healthy plants. Palisade cells short, chloroplasts smaller and fewer than nor- mal. Transmission : By needle puncture into rhizomes immersed in filtered juice of dis- eased plant. By contamination of soil with fragments of diseased leaves or rhizomes. Literature: Klebahn, Bericht. d. Deutsch. Bot. Gesellsch., 15, 1897, 527- 536; Ztschr. wissensch. Biol., Abt. E, Planta, 1, 1926, 419-440; 6, 1928, 40-95; Phytopath. Ztschr., 4, 1932, 1-36; 9, 1936, 357-370. 4. Galla verrucae Blodgett. (Phyto- path., S3, 1943, 30.) From Latin verruca, wart. Originally spelled verruca, ap- parently by a typographical error, which was corrected twice on the following page, once in a statement that the genitive ver- rucae had been given as specific epithet. Common name : Peach-wart virus. Host : ROSACEAE — Primus persica (L.) Batsch, peach. Geographical distribution : United States (Idaho, Washington, Oregon). Induced disease : In peach, no charac- teristic effect on foliage. Fruits blis- tered, welted, later with warty out- growths conspicuously raised. Affected tissues light tan to red, rough, cracked and russeted, or smooth. Gumming usual, often severe. Warty tissue super- ficial; underlying tissues coarse, filled with gum pockets, but not abnormal in flavor. Warty tissue may be hard and bony, but more often it is merely tougher than normal. Transmission : By budding and in- arching. Literature: Blodgett, Phytopath., 31, 1941, 859-860 (Abst.) ; 33, 1943, 21-32. 5. Galla zeae McKinney. (Jour. Washington Acad. Sci., 34, 1944, 328.) From Latin sea, a kind of grain. Common name : Wallaby-ear disease virus. Host: GRAMINEAE—Zea mays L., corn (maize). Geographical distribution : South- eastern Queensland, Australia. Induced disease: In corn (maize), small swellings on secondary veins on undersides of young leaves, spreading to base and tip of leaf along veins; plant dwarfed, becoming abnormall}' deep green and deficient in development of pollen; silk, cobs, and grain retarded in growth. FAMILY CHLOROGENACEAE 1159 Transmission : By leafhopper, Cicadula himacidata Evans (CICADELLIDAE). Literature: Schindler, Jour. Austral. Inst. Agr. Sci., 8, 1942, 35-37. Genus VI. Fractilinea McKinney. (Jour. Washington Acad. Sci., 34, 1944, 148.) Viruses of the Stripe-Disease Group ; hosts grasses ; insect vectors, cicadellid and fulgorid leafhoppers. Generic name from Latin, meaning interrupted and line. Tlie type species is Fractilinea maidis (Holmes) McKinney. Key to the species of genus Fractilinea. I. Vectors, cicadellid leafhoppers. 1. Fractilinea maidis. 2. Fractilinea oryzae. 3. Fractilinea tritici. 4. Fractilinea quarta. II. Vectors, fulgorid leafhoppers. 5. Fractilinea zeae. 6. Fractilinea avcnae. 1. Fractilinea maidis (Holmes) Mc- Kinney. {M armor maidis Holmes, Handb. Phytopath. Viruses, 1939, 56; Fractilinea maidis JMcKinney, Jour. Washington Acad. Sci., 34, 1944, 149.) From New Latin mays, corn (i.e. maize). Common name: Maize-streak virus. Hosts: GRAMINEAE—Zea mays L., corn (maize); Digitaria horizontalis Willd., Eleusine indica Gaert.; Sac- charum officinarum L., sugar cane. Geographical distribution : Africa. Induced disease : In corn, pale spots at base of young leaf, followed by chlorotic spotting and streaking of subsequently formed leaves. Virus moves rapidly (up to 40 cm in 2 hours at 30° C) after intro- duction into host plant by insect. More virus in chlorotic spots than in green areas of affected leaves. Transmission : B3' leafhoppers, Cica- dulina (= Balclutha) mbila (Naude), C. zeae China, and C. storeyi China (CICA- DELLIDAE). In C. mbila ability to transmit this virus is controlled by a sex-linked dominant gene ; active male (AX) (Y), inactive male (aX) (Y), inac- tive female (aX) (aX), active female (AX) (AX) or (AX) (aX). Inactive in- dividuals ingest virus when feeding, but can become infective only if the intestine is wounded purposely or accidentally. If inoculated artificially by introducing virus into blood, both active and inactive insects become infective. Incubation period, 6 to 12 hours at 30° C. Young not infected through the egg. Infective leaf- hopper cannot transmit virus unless feed- ing puncture exceeds a minimum period, about 5 minutes in duration. This virus has not been transmitted to its plant hosts by inoculation of expressed juices. Filterability : At pH 6, passes Cham- berland Li and L3, Berkefeld V and N, filters; retained by Seitz E K filter disc. Literature: Storey, Ann. Appl. Biol., 12, 1925, 422-439; 75, 1928, 1-25; 19, 1932, 1-5; Proc. Roy. Soc, B, 112, 1932, 46-60; 113, 1933, 463-185; 125, 1938, 455-477; Ann. Appl. Biol., 21, 1934, 588-589; 24, 1937, 87-94; East Afr. Agr. Jour., 1, 1936, 471-475 ; Storey and McClean, Ann. Appl. Biol., 17, 1930, 691-719. Strains : Two strains that differ radi- cally from the type, var. typicum H. {loc. cit., 56), have been given varietal names, as follows : la. Fractilinea maidis var. sacchari H. (loc. cit., 57). From New Latin Sac- charum, generic name of sugar cane. Common name : Cane -streak strain of 1160 MANUAL OF DETERMINATIVE BACTERIOLOGY maize-streak virus. Differing from the type strain in being specialized for at- tacking sugar cane, in which the type (from maize) tends to be localized or finally lost with resultant spontaneous recovery of the temporary host. The cane -streak strain usually spreads readily in the cane plant; leaves become much marked with broken, narrow, pale, longi- tudinal stripes and spots ; stems remain unaffected. One variety of sugar cane, P.O.J. 213, is resistant and, if infected, tends to recover. (McClean, Intern. Soc. Sugar Cane Techn., Bull. 27, 1932; Proc. So. Afr. Sugar Techn. Assoc, 1936, 1-11; Storey, Rept. Imp. Bot. Conf., London, 1924, 132-144; Union So. Afr. Dept. Agr., Sci. Bull. 39, 1935; Ann. Appl. Biol., 11, 1930, 691-719. lb. Fractilinea maidis var. mitia H. {loc. cit., 58). From Latin mitis, mild. Common name : Mottle strain of maize - streak virus. Differing from the typical strain by the mildness of the disease in- duced in corn (maize), transitory chlo- rotic mottling of newly developed leaves, followed by fading of mottling and pro- duction of apparently normal leaves. Young leaves, while mottled, are less rigid tlian normal and may not remain as nearly erect as healthy leaves. (Storey, Ann. Appl. Biol., 2^, 1937, 87-94.) 2. Fractilinea oryzae (Holmes) comb, nov. {Marmor oryzae Holmes, loc. cit., 64.) From Latin oryza, rice. Common name : Rice dwarf -disease virus. Hosts: GRAM I NEAE— Oryza saliva L., rice. Experimentally, also Alope- cur^ts fulvxis L.; Avena saliva L., oat; Echinochloa crusgalli Beauv. var. edulis Honda; Panicinn miliaceum L.; Poa pralensis L.; Secale cereale L., rye; Trilicum vulgar e Vill., wheat. Insusceptible species : GRAMINEAE — Zea mays L., corn (maize); Hordeum vulgareh., harley; Selaria italica Beauv., foxtail millet; Andropogon sorghum Brot. (= Holcus sorghum L.), sorghum. Geographical distribution : Japan, Phil- ippine Islands. Induced disease: In rice, yellowish green spots along veins of young leaf, fol- lowed by chlorotic spotting and streaking of subsequently formed leaves. Growth stunted, internodes and roots abnormally short, forming a dwarf plant. Vacuolate intracellular bodies, 3 to 10 by 2.5 to 8.5 microns in size, close to nuclei of cells in affected tissues. Transmission : By leaf hoppers, Nepho- tetlix apicalis var. cincticeps *Uhler, N. bipunctalus Fabr., and Dellocephalus dor- saZts Motsch. (CICADELLIDAE). Vi- rus transmitted through some of the eggs but through none of the sperm of infected individuals of N. apicalis. Transfer from individuals thus infected through the egg to their progeny likewise demon- strated, even to the 7th generation. This is the only confirmed instance of trans- mission of a phytopathogenic virus through the eggs of an insect vector and is considered as evidence that the virus multiplies within the body of its vector as well as in its plant hosts. Incubation period in insect usually 30 to 45 days after first feeding on an infected plant, some- times as short as 10 or as long as 73 days ; nymphs from viruliferous eggs do not be- come infective until 7 to 38 (average 19) days after emergence. Transmission by inoculation of expressed juice has not been demonstrated. No transmission through seeds from diseased rice plants. No soil transmission. Literature: Agati et al., Philippine Jour. Agr., 12, 1941, 197-210; Fukushi, Trans. Sapporo Nat. Hist. Soc, 12, 1931, 35-41; Proc. Imp. Acad., Tokyo, 9, 1933, 457-460; Jour. Fac. Agr. Hokkaido Imp. Univ., 37, 1934, 41-164; Trans. Sapporo Nat. Hist. Soc, 13, 1934, 162-166; Proc. Imp. Acad., Tokyo, 11, 1935, 301-303; 13, 1937, 328-331; 15, 1939, 142-145; Jour. Fac Agr. Hokkaido Imp. Univ., 45, 1940, 83-154; Katsura, Phytopath., FAMILY CHLOROGEXACEAE 1161 S6, 1936, 887-895; Takata, Jour. Japan Agr. Soc, 171, 1895, 1-4; 172, 1896, 13-32 (Takata's papers, in Japanese, constitute the first published record of transmission by an insect of a virus causing disease in a plant, the leaf hopper Deltocephaliis dor- salis Motsch. transmitting dwarf -disease virus to rice; see Fukushi, 1937, cited above ) . 3. Fractilinea tritici^IcKinney. (Jour. Washington Acad Sci., 34, 1944, 327.) From Latin triticum, wheat. Common name : Winter-wheat mosaic virus. Hosts: GRAM IN EAE— Triticum aes- tivum L., wheat; Secale cereale L., rye; Avena byzaniina; A. fatua L., wild oat; A. saliva L., oat; Hordeum vulgare L., barley. Geographical distribution: Union of Soviet Socialist Republics. Induced disease : In winter wheat and rye, chlorotic mottling; profuse branch- ing. In winter wheat, phloem necrosis; chloroplasts few, small; vacuolate inclu- sions in cells; nuclei enlarged and with extra nucleoli ; no protein crystals of the pupation-disease type in affected cells. In spring wheat, barley, and oats, chlo- rotic mottling without profuse branching ; no proliferation of flowers, but grain is rarely formed, most infected plants dying before this stage of growth. Transmission : By leafhopper, Delto- cephalus striatus L. (CICADELLIDAE), with incubation period of 15 to 18 days. Not by inoculation of expressed juice. Not through soil. Literature : Zazhurilo and Sitnikova, Compt. rend. Acad. Sci. U. R. S. S., ^5, 1939, 798-801 ; 26, 1940, 474-478; 29, 1940, 429-432; Proc. Lenin Acad. Agr. Sci., U. R. S. S., 6, 1941, 27-29. [Rev. Appl. Myc, 19, 1940, 268; 20, 1941, 157, 396; 22, 1943, 59]. 4. Fractilinea quarta (Holmes) cotrib. nov. (Marmor quartian Holmes, loc. cii., 65.) From Latin quarius, fourth. Common name : Sugar-cane chlorotic- streak virus or fourth -disease virus. Host : GRAMINEAE—Saccharum offi- cinarum L., sugar cane. Geographical distribution : Java, Queensland, Hawaii, Puerto Rico, Colom- bia, United States (Louisiana). Induced disease : In sugar cane, reduc- tion of growth rate; wilting at midday; long, narrow, longitudinal streaks, of creamy or white color, in the leaves. Streaks 1/16 to 3/16 inches wide, gen- erally less than 1 foot long, fragmenting. Transmission : Bj^ leafhopper, Draecu- lacephala poriola Bsdl (CICADELLI- DAE). Not demonstrated by inocula- tion of expressed juice. Thermal inactivation: In cuttings, at 52° C in less than 20 minutes. Literature: Abbott, Phytopath., 28, 1938, 855-857; Sugar Bull., 16, 1938, 3-4; Abbott and Ingram, Phytopath., 3^, 1942, 99-100; Bell, Queensland Agr. Jour., 40, 1933, 460-464 ; Martin, Hawaiian Planters' Rec, 34, 1930, 375-378; Hawaiian Sugar Planters' Assoc. Proc, 53, 1934, 24-35. 5. Fractilinea zeae (Holmes) comb, nov. {Marmor zeae Holmes, loc. cit., 59.) From New Latin Zea, generic name for corn (maize), from Latin zea, a kind of grain. Common name: Maize-stripe virus. Host: GRAMINEAE~Zea mays L., corn (maize). Insusceptible species: GRAMINEAE — Saccharum officinarum L., sugar cane. Geographical distribution: Hawaii, Tanganyika, Mauritius, Trinidad, Cuba. Not in United States. Induced disease: In corn (maize), at first few, elongated, chlorotic lesions near base of young leaf, later enlarging and fus- ing to form continuous stripes. Subse- quently formed leaves banded and striped variously. Vacuolate intracellular in- clusions in cells of affected areas. Transmission : By leafhopper, Pere- grinus maidis (Ashm.) (F ULGORI- DAE) ; the incubation period in this in- 1162 MANUAL OF DETERMINATIVE BACTERIOLOGY sect is usually between 11 and 29 days, although shorter periods have been dem- onstrated in a few cases. Virus may per- sist in the insect host until death, but may become exhausted earlier. Not by aphid, Aphis maidis Fitch (APIIIDI- DAE). Not by inoculation of expressed juice. Literature: Briton-Jones, Trop. Agr., 10, 1933, 119-122; Carter, Ann. Ent. Soc. Am., 34, 1941, 551-556; Kunkel, Bull. Hawaiian Sugar Planters' Assoc, Bot. Ser., 3, 1921, 44-57; 1924, 108-114; Hawaiian Planters' Rec, 26, 1922, 58-64; Phytopath., 17, 1927, 41 (Abst.); Stahl, Trop. PI. Res. Found., Bull. 7, 1927; Storey, Rept. of Plant Pathologist, Amani Agr. Res. Station, 4th Ann. Rept., 1931-32, pp. 8-13. 6. Fractilinea avenae McKinney. (Jour. Washington Acad. Sci., 34, 1944, 327.) From Latin avena, oats. Common name : Pupation-disease virus. Hosts: GRAM I NEAE— Avena saliva L., oat; Triticuvi aesiivum L., wheat; Echinochloa crusgalli Beauv.; Setaria viridis; rarely, Agropyron repens (L.) Beauv. and Bromus inerviis Leyss. Ex- perimentally, also Hordeum vulgare L., barley; Panicum miliaceum L., millet; Oryza sativa L., rice; Secale cereale L., rye; Zea mays L., corn (maize). Geographical distribution : West Si- beria. Induced disease- In oat, chlorotic mottling, profuse development of shoots, proliferation pf flowers with change to leaf-like structures. Protein crystals in affected cells have been regarded as accumulated virus. Transmission : By leaf hopper, Delphax striatella Fallan (FULGORIDAE), es- pecially first and second instar nymphs; fifth instar nearly immune to infection. Incubation period, 6 days or more. Virus overwinters in insect as well as in plants. Not transmitted from an infected leaf- hopper to its progeny. Not through soil. Not through seeds from infected plants. Literature : Sukhov et al., Compt. rend. Acad. Sci., U. R. S. S., 30, 1938, 745-748; 26, 1940, 479-482, 483-486. FAMILY MARMORACEAE 1163 FAMILY II. MARMORACEAE HOLMES EMEND. (Handb. Phytopath. Viruses, 1939, 16.) Viruses of the ^losaic Group, inducing diseases usually characterized b}' persistent chlorotic or necrotic spotting, and often by mottling. The family is here extended to include several small groups of viruses, formerly assigned independent family rank, but sharing a tendency to aphid transmission, so far as known, and inducing diseases characterized bj- abnormal growth habit, thickening and rolling of leaves, or dwarfing, traits not incompatible with the characters of the present group. Should anj' one of these small groups become the center of a large assemblage of new viruses in the future, separate familial status for it might again be advantageous. In the combined group- ing here used, specific vectors, so far as known, are aphids (APHIDIDAE). Key to the genera of Jamily Marmoraceae. I. Viruses of the Typical Mosaic-Disease Group. Genus I. Marmor, p. 1163. II. Viruses of the Spindle -Tuber Group. Genus II. Acrogenus, p. 1202. III. Viruses of the Leaf -Roll Group. Genus III. Corium, p. 1203. IV. Viruses of the Dwarf -Disease Group. Genus IV. Nanus, p. 120G. V. Viruses of the Rough-Bark Group. Genus V. Rimocortius , p. 1208. VI. Viruses of the Symptomless Group. Genus VI. Adelonosus , p. 1211. Genus I. Marmor Holmes. {Loc. cit., 16) Viruses inducing t3'^pical mosaic diseases in various plants. Generic name from Latin marmor, a mottled substance. The type species is Marmor tabaci Holmes. Key to the groups within germs Marmor. A. Relatively resistant to heat inactivation, usually requiring more than 10 minutes at 85 to 90° C for complete inactivation. 1. Tobacco -IMosaic Virus Group. B. Relatively susceptible to heat inactivation, requiring less than 10 minutes at 85 to 90° C for complete inactivation. a. Replacing potato-veinbanding virus in mixed infections. 2. Tobacco-Etch Virus Group, aa. Not replacing potato-veinbanding virus in mixed infections. 3. Cucumber-Mosaic Virus Group. C. Many additional species cannot yet be grouped into definite subdivisions of the genus ; they constitute a residual or 4. Miscellaneous Mosaic-Virus Group. Key to the species of the Tobacco-Mosaic Virus Group. Viruses relatively resistant to heat inactivation, requiring in most cases more than 10 minutes at 85 to 90° C for complete inactivation. Insect vectors as yet unknown under natural conditions. 1164 MANUAL OF DETERMINATIVE BACTERIOLOGY I. Found in nature principally in solanaceous plants; Cucurbitaceae insus- ceptible. Chlorotic mottling in some hosts, necrotic lesions in others as result of experimental infection. Suspensions show anisotropy of flow. 1. M armor tabaci. 2. Marmor constans. II. Found in nature only in cucurbitaceous plants; Solanaceae insusceptible. Only mottling as result of experimental infection. Suspensions show marked anisotropy of flow. 3. Marmor astrictum. III. Found only in leguminous plants. Chlorotic lesions in some varieties of the common snap -bean plant, necrotic lesions in others, as a result of experi- mental infection. 4. Marmor laesiofaciens . IV. Found in greenhouses confined to roots and lower parts of plants. Only necrotic lesions as result of experimental infection. Suspensions do not show anisotropy of flow. 5. Marmor lethale. V. Found in tomato and experimentally transmissible to a number of species of plants in this and other families. Resembling the preceding in a number of physical characteristics, including failure to show anisotropy of flow. 5a. Marmor dodecahedron. 1. Marmor tabaci Holmes. (Holmes, Handb. Phytopath. Viruses, 1939, 17; Musivum tabaci Valleau, Phytopath., 30, 1940, 822; Fhytovirus nicomosaicum Thornberry, Phytopath., 31, 1941, 23.) From New Latin Tabacum, early generic name for tobacco. Common names : Tobacco-mosaic virus, tomato-mosaic virus. Hosts : SOLANACEAE— Nicotiana tabacum, L., tobacco; Lycopersicon escu- lentum Mill., tomato; and Capsicum frutescens L., garden pepper, among crop plants; nearly all, if not all, solanaceous plants can be infected, although in some the virus remains localized at or near the site of inoculation. PLANTAGINA- CEAE — A strain of this virus has been found in nature infecting Plantago lance- olata L., ribgrass, P. major L. and P. rugelii Dene., common broad-leaved plantains. Experimental hosts are widely distributed through many related families of plants. Geographical distribution : World-wide. Induced disease : In most varieties of tobacco, yellowish-green primary lesions, followed by clearing of veins, distortion and greenish-yellow mottling of newly formed leaves. In Ambalema tobacco, no symptoms, virus being restricted to inoculated leaves or those nearby. Strains of tobacco showing necrotic ef- fects have been produced recently. In tomato, no obvious primary lesions, sys- temic disease characterized by greenish- yellow mottling of foliage, moderate dis- tortion of leaf shape, and a reduction of fruit yield not exceeding 50 per cent. If some strain of potato-mottle virus (Mar- mor dubiinn) is also present, a more severe disease is induced; this is known as double-virus streak, and is characterized by systemic necrosis. In most varieties of garden pepper, yellowish primary lesions followed by systemic chlorotic mottling. In the Tabasco pepper and its recent derivatives, recovery by abscission of inoculated leaf, after localization of virus in necrotic primary lesions. Vacu- olate intracellular inclusions are found in chlorotic tissues of all hosts that show distinct chlorotic mottling. Transmission : By slight abrasive con- tacts. By inoculation of expressed juice. To some extent by the aphids, Myzus pseudosolani Theob., M. circumflexus (Buckt . ) , Macrosiphum solanifolii Ashm . , and Myzus persicae (Sulz.) {APHIDI- DAE). By grafting. Through soil. FAMILY MARMORACEAE 1165 Through dodder, Cuscuta campestris Yuncker (CONVOLVULACEAE), with- out infecting this plant vector. Not through pollen from diseased plants. Not through seeds from diseased tobacco ; seed transmission has been reported in the case of recently ripened seeds from diseased tomato. Serological relationships : Precipitin test gives cross reactions between all known strains, except those characterized by failure to spread systemicallj' in to- bacco. No cross reactions with other viruses except weakly with cucurbit - mosaic virus {M armor astrictum). Type and other strains of tobacco-mosaic virus give cross reactions in complement -fixa- tion and neutralization tests. Immunological relationships : Plant protection tests, particularly in Nicotiana sylvestris Spegaz. and Comes, have dem- onstrated that tissues invaded by any strain of this virus are immune to subse- quent infection by the tomato aucuba- mosaic strain of tobacco-mosaic virus, indicating a group relationship not shared by other viruses, such as cucumber- mosaic virus or tobacco-ringspot virus. Thermal inactivation : At 88 to 93° C in 10 minutes ;at 86 to 92° C in 30 minutes. Filterability : Tobacco -mosaic virus was the first virus shown to be filterable, by Iwanowski in 1892; its filterability was confirmed and interpreted by Beijerinck in 1898. Other properties: The ultimate parti- cles of tobacco-mosaic virus have been shown to be rod-shaped and isotropic, sometimes associated in pairs, end to end. Under proper conditions, thread-like paracrystals are formed. Specific gravity has been determined as about 1.37, re- fractive index as about 1.6. Lsoelectric point between pH 3.2 and 3.5. Suspen- sions in media of lower refractive indices show anisotropy of flow. Sedimentation constants, at 20° C, 187 X IQ-i^ cm per sec. per dyne at infinite dilution for un- aggregated virus and 216 X 10"^^ cm per sec. per dyne for associated particles. The computed average length of the virus unit is about 272 millimicrons; diameter, 13.8 millimicrons. Electron micrographs show that characteristic particles are rod- like, between 10 and 20 millimicrons in width, variable in length, but in some preparations averaging 270 millimicrons in length for single units, 405 to 540 milli- microns in length for associated pairs; X-ray measurements in air-dr}^ gel show- width 15.20 ± 0.05 millimicrons. Solu- tions stronger than about 1 .3 per cent sep- arate into layers, the lower spontaneously doubly refracting and more concentrated than the upper. At concentrations of electrolytes somewhat less than are re- quired to precipitate the virus as fibres or needle-shaped paracrystals, the solu- tions form clear gels that become fluid on shaking or diluting (at pH 7 and 30° C). The virus is destroyed by high-fre- quenc}- sound radiation, by pressures be- tween 6000 and 8000 kilograms per square centimeter, and by hydrogen-ion con- centrations above pH 11 or below pH 1. It is relatively stable between pH 2 and pH 8. It is rapidly broken down in 6 M urea solutions, in the presence of salts, to low-molecular-weight compo- nents devoidof activity. Analysisof puri- fied virus : carbon 47.7 per cent, hydrogen 7.35 per cent, nitrogen 15.9 per cent, sul- fur 0.24 per cent, phosphorus 0.60 per cent, lipoid 0.0 per cent, carbohydrate 1.6 to 2.0 per cent . A revised estimate of the sulfur content is 0.20 per cent, probably all in cysteine ; no methionine has been detected in the typical variety of this virus. The percentages of the following substances in the virus are: tyrosine 3.9, tryptophane 4.5, proline 4.6, arginine 9.0, phenylalanine 6.0, serine 6.4, threonine 5.3, cysteine 0.68, alanine 2.4, aspartic acid 2.6, glutamic acid 5.3, leucine 6.1, valine 3.9, nucleic acid 5.8, and amide nitro- gen 1.9, collectively accounting for about 68 per cent of the total weight. Virus formation ceases in infected host tissues immersed in 0.0002 molar sodium cyanide solution, beginning again after removal of cvanide. 1166 MANUAL OF DETERMINATIVE BACTERIOLOGY Literature : The literature dealing with tobacco-mosaic virus is too voluminous to permit citation of more than a few repre- sentative publications. Allard, U. S. Dept. Agr., Bull. 40, 1914; Bawden and Pirie, Proc. Roy. Soc. London, Ser. B., 123, 1937, 274-320; Beale, Jour. Exp. Med., 54, 1931, 463-473; Beijerinck, Ver- handel. Konink. Akad. Wetenschappen te Amsterdam, II, 6, 1898, 3-22; Grant, Phytopath., U, 1934, 311-336; Hoggan, Jour. Agr. Res., 49, 1934, 1135-1142; Iwanowski, Bull. Acad. Imp. Sci. St. Petersburg, Ser. 4, 3, 1892, 67-70; Jensen, Phytopath., 23, 1933, 964-974; Johnson, Science, 64, 1926, 210; Kausche et al., Naturwiss., S7, 1939, 292-299; Knight, Jour. Biol. Chem., 147, 1943, 663-666; Kunkel, Phytopath., 24, 1934, 437-466; Lauffer, Jour. Am. Chem. Soc, 66, 1944, 1188-1194; Price, Phytopath., 23, 1933, 749-769; Stanley, Phytopath., 26, 1936, 305-320; Takahashi and Rawlins, Proc. Soc. Exp. Biol, and Med., 30, 1932, 155- 157 ; Valleau and Johnson, Kentucky Agr. Exp. Sta., Bull. 376, 1937; Vinson, Sci- ence, 66, 1927, 357-358; Woods, Science, 91, 1940, 295-296. Strains : A great number of variant strains have been isolated both experi- mentally and from plants infected in nature. These usually share with the type variety most of the fundamental properties, particle size, especially width, stability at relatively high temperatures, longevity in storage, some common anti- gens. The following have been distin- guished from the type, var. vulgar e H. {loc. cit., 17), by varietal names : la. Marmor tabaci var. aucuba H. {loc. cit., 20). A group of isolates producing necrotic local lesions in inoculated leaves of Nicotiana sylvestris Spegaz. and Comes ; useful in identifying many other strains of this virus which on prior ap- plication protect the tissues of this plant from the necrotic effects of aucuba-type strains (Smith, Ann. Appl. Biol., 18, 1931, 471-493; Kunkel, Phytopath., 24, 1934, 437-466). lb. Marmor tabaci var. deformans H. (loc. cit., 22). Producing exceptionally severe malformation of tomato foliage. (Ainsworth, Ann. Appl. Biol., 24, 1937, 545-556). Ic. Marmor tabaci var. canadense H. (loc. cit., 23). Producing a necrotic type of streak disease in tomatoes (Jarrett, Ann. Appl. Biol., 17, 1930, 248-259). Id. Marmor tabaci var. lethale H. {loc. cit., 24). Producing spreading necrotic lesions in tobacco and tomato under ex- perimental conditions (Jensen, Phyto- path., 27, 1937, 69-84 ; Norval, Phytopath. 28, 1938, 675-692). le. Marmor tabaci var. plantaginis H. (Phytopath., 31, 1941, 1097). Specially adapted in nature for systemic spread in species of Plantago. This variety con- tains histidine (0.55 per cent) and methi- onine (2 per cent) not found in the type of the species. If. Marmor tabaci var. obscurum H. (Handb. Phytopath. Viruses, 1939, 25). Systemic in tobacco without producing obvious disease under experimental con- ditions (Holmes, Phytopath., 24, 1934, 845-873 ; 26, 1936, 896-904 ; Jensen, Phyto- path., ^7, 1937,69-84). Ig. Marmor tabaci var. immobile H. {loc. cit., 26). Produces chlorotic pri- mary lesions in experimentally infected tobacco, but rarely becomes systemic. (Jensen, Phytopath., 23, 1933, 964-977; 27, 1937,69-84). Ih. Marmor tabaci var. artuin H. {loc. cit., 27). Necrotic lesions experiment- ally induced in Nicotiana glutinosa L. {SOLAN ACE AE) are much smaller than those of the type variety (Jensen, Phy- topath., 27, 1937, 69-84). li. Marmor tabaci var. siccans Doolittle and Beecher. (Phytopath., 32, 1942, 991). Causing necrosis and shriveling of tomato foliage. FAMILY MARMORACEAE 1167 2. Marmor constans McKinney. (Jour. Washington Acad. Sci., 34, 1044, 326.) From Latin constans, fixed. Common name : Tobacco niild dark- green mosaic virus. Hosts : SOLA NACEAE—Nicotiana glauca R. Grah., tree tobacco. Insusceptible species: SOLAN A- CEAE — Lycopersicon esculentum Mill., tomato. CUCURBITACEAE— Cucumis sativus L., cucumber. Geographical distribution : Islands of Grand Canary and Teneriffe. Induced disease : In Nicotiana glauca, sj'stemic chlorotic mottling. Transmission : By inoculation of ex- pressed juice. No insect vector is known. Thermal inactivation: At about 86°C. in 10 minutes. Literature : McKinney, Jour. Agr. Res., 39, 1929, 557-578; Am. Jour. Bot., 28, 1941, 770-778; Peterson and McKinney, Phytopath., 28, 1938, 329- 342; Thornberry and McKinney, ibid., 29, 1939, 250-260. 3. Manner astrictum Holmes. (Holmes, Handb. Phytopath. Viruses, 1939, 27; Musivum astrictum Valleau, Phytopath., SO, 1940, 823.) From Latin astrictus, limited, in reference to host range. Common names : Cucurbit-mosaic virus, English cucumber-mosaic virus. Hosts: CUCURB IT ACE AE— Cucu- mis sativ2(s L., cucumber; C. anguria L., gherkin ; C. melo L., melon ; Citrullus vul- garis Schrad., watermelon; only cucur- bitaceous plants have appeared to be susceptible thus far. Insusceptible species : All tested sol- anaceous species. CUCURBITA- CEAE — Bryonia dioica L.; Cucurhita pepo L., vegetable marrow. LEGU- MINOSAE — Phaseolus vulgaris L. var. Golden Cluster. Geographical distribution : England. Induced disease : In cucumber, clearing of veins and crumpling in young leaves, followed by a green-mosaic mottling, with blistering and distortion of newly formed leaves. Plant stunted. Fruit un- marked or slightly mottled. Diseased plants less obviously affected during win- ter months. Transmission : By inoculation of ex- pressed juice. No insect vector is known . Serological relationships : Weak cross precipitin reactions and full cross-neu- tralization reactions with tobacco -mosaic virus (Marmor tabaci). Two common antigens postulated. Preparations of vi- rus that have been inactivated by treat- ment with nitrous acid or X-raj^s are still antigenic. Thermal inactivation : At 80 to 90° C in 10 minutes. Filterability : Passes Pasteur-Cham- berland filters Lj to L?, and membranes of 150 millimicrons average pore diameter. Other properties : Virus, infectious in dilution of 10"^°, is present to the extent of 0.2 to 0.3 gram per liter of juice from diseased plants. Preparations show sheen and anisotropy of flow, indicating rod-shaped particles. Solutions stronger than 2.5 per cent separate into 2 layers at room temperature, the lower being the more concentrated and birefringent. Precipitates with ammonium sulfate show needle-shaped paracrystals. Sedimenta- tion constants Sjo" = 173 X lO'i^ cm. sec.~J dyne~^ and about 200 X 10"!^ cm. sec.~i dyne~i. Virus withstands drying without inactivation but with partial loss of ability to show anisotropy of flow and with reduction of serological activity to about half. Tryptophane content 1.4 per cent, phenylalanine 10.2 per cent, the first lower and the second higher than in tobacco-mosaic virus. Literature: Ainsworth, Ann. Appl. Biol., 22, 1935, 55-67; Bawden and Pirie, Nature, 139, 1937, 546-547; Brit. Jour. Exp. Path., 18, 1937, 275-291; Knight, Arch. Virusf., 2, 1942, 260-267; Knight and Stanley, Jour. Biol. Chem., I4I, 1941, 29-38; 141, 1941, 39-49; Price, Am. Jour. Bot., 27, 1940, 530-541; Price and Wyc- koff, Nature, 14I, 1938, 685. 1168 MANUAL OF DETERMINATIVE BACTERIOLOGY Strains : A distinctive strain has been distinguished from the type, var. chloro- genus H. {loc. cit., 27), by the varietal name : 3a. Marmor astrictum var. aucuba H. (loc. cit., 29). Differing from the type of this species by inducing a yellow-mottl- ing, rather than a green-mottling, mosaic in cucumber (Ainsworth, Ann. Appl. Biol., 32, 1935, 55-67). 4. Marmor laesiofaciens Zaumeyer and Harter. (Jour. Agr. Res., 67, 1943, 305.) From Latin laesio, substantive from laedere, to injure, and participle from facere, to make. Common name: Bean-mosaic virus 4; southern bean mosaic virus 1 . Hosts : LEG UMINOSAE—Phaseolus vulgaris L., bean. Experimentally, also Phaseolus lunatus L., sieva bean; Soja max Piper var. Vii-ginia, Virginia soy bean . Insusceptible species : All tested spe- cies in families other than the LEGU- MINOSAE. Geographical distribution: United States (Louisiana). Induced disease : In bean, systemic chlorotic mottling in some varieties, localized necrosis in others; in a few varieties, systemic necrosis. In mot- tling-type varieties, chlorotic mottling of foliage; pods marked by dark green blotches or shiny areas, slightly mal- formed, short, frequently curled at end. In necrotic-type varieties with localized response, bearing a dominant gene lacking in mottling-type varieties, reddish ne- crotic lesions at the site of inoculation ; no evidence of systemic spread of virus. In varieties showing systemic necrosis, pin- point or slightly larger necrotic primary lesions with veinal necrosis of inoculated leaf; systemic veinal necrosis, distortion and curling of affected leaves, drooping at the pulvini ; stem and petiole streak ; eventual death of plant . Transmission : By inoculation of ex- pressed juice. Through seeds from in- fected plants. Serological relationships : Not demon- strated. Immunological relationships : Previous infection with bean-mosaic virus, Marmor phaseoli, does not protect against infec- tion with this virus. Thermal inactivation : At 90 to 95° C, time not stated, probably 10 minutes. Other properties : Withstands dilution of 1 : 500,000 and aging 32 weeks at 18° C. Literature : Zaumeyer and Harter, Phy- topath., 32, 1942, 438-439; 33, 1943, 16; 34, 1944, 510-512; Jour. Agr. Res., 67, 1943, 295-300, 305-328. Strains : A strain differing from the type lias been given the varietal name : 4a. Marmor laesiofaciens var. ininus Zaumeyer and Harter. (Jour. Agr. Res., 67 , 1943, 305.) From Latin minor, lesser. Differing from the type by inducing formation of slightly less diffuse and spreading lesions in necrotic-type bean leaves; also by inducing milder early symptoms and more severe late symptoms in mottling-type beans. Passes through seeds from infected plants to infect seed- lings grown from them. Found in addi- tional localities in the United States (California, Colorado, Idaho, Maryland). 5. Marmor lethale H. (loc. cit., 86). From Latin lethalis, causing death. Common name : Tobacco-necrosis virus. Hosts: SOLAN ACE AE—Nicotiana tabacum L., tobacco; A'', gluiinosa L.; N. langsdorffii Weinm. ; Lycopersicon escu- lenium Mill., tomato; Solanum nigrum L. COMPOSITAE—k^iQx. GE RANI- ACE AE — Pelargonium hortorum Bailey. LEG UM I NOSAE— Phaseolus vulgaris L., bean. Confined to roots of these natural hosts except in the cases of Nicotiana tabacum and N . gliitinosa in which lower leaves are sometimes in- vaded; necrotic lesions along midrib and larger veins in these. No obvious mani" festations of disease in infected roots. Experimentally to plants in many fami- lies with production of localized necrotic lesions only. FAMILY MARMORACEAE 1169 Geographical distribution: England, Scotland, Australia. This virus has been found only in greenhouses. Induced disease : In tobacco, necrosis of midrib and larger veins of first-de- veloped pair of leaves, between Novem^ ber and February. Virus also in roots of many healthy -looking plants through- out the j'ear. Upon artificial inoculation of foliage, numerous small brown necrotic local lesions are produced. Yield of virus from infected plant 0.02 mg per cc of ex- pressed juice, on the average. Transmission : By contamination of soil with virus. No insect vector is known. Experimentally, by inoculation of ex- pressed juice. Serological reactions : Precipitates with homologous antiserum. No cross reac- tion with tomato bush5^-stunt or tobacco- mosaic viruses. Immunological relationships : Protec- tion tests show lack of relationship to tobacco-mosaic virus, tobacco-ringspot virus, tomato-ringspot virus, cucumber- mosaic virus, and the severe-etch strain of tobacco-etch virus. Thermal inactivation : At 90 to 92° C in 10 minutes. Filterability : Average particle diame- ter 20 to 30 millimicrons as determined by filtration through Gradocol membranes; other reports give diameter as 13 to 20 millimicrons by filtration (14 to 19 milli- microns by radiation experiments, about 20 millimicrons from electron micro- graphs). Other properties : Infectious after stor- age for months in dried leaves and after storage for half a j'ear in absolute ethyl alcohol at room temperature. Specific gravity 1.3. More soluble in ammonium sulfate solutions at 0° C than at room temperature. Composition : Carbon 44.8 to 45.3 per cent, nitrogen 15.5 to 16.5 per cent, hydrogen 6.5 to 7.0 per cent, phos- phorus 1.4 to 1.7 per cent, sulfur 1.1 to 2.0 per cent, carbohydrate 7.0 to 9.0 per cent ; ash 5.8 to 7.0 per cent (3 to 5 per cent after prolonged dialysis at pH 3). Nucleic acid of the ribose type has been isolated. No anisotropy of flow in solu- tion but crj'stals are birefringent, show- ing sharp extinctions parallel to, and at right angles to, the plane of the crj^stal when examined edge-on in a polarizing microscope. Sedimentation constant, S2o° = 112 X 10~i'; in other preparations a crystalline component with sedimenta- tion constant 130 X 10"^' and an amor- . phous component with sedimentation constant 58 X lO^^^ have been reported, as well as small amounts of a substance with sedimentation constant 220 X 10~^^ Strains : Isolates of tobacco-necrosis virus serologically distinct but not other- wise different from each other appear to impl.y the existence of several strains of this virus, or of a closely related group of viruses, in England. Literature: Bawden, Brit. Jour. Exp. Path., 22, 1941, 59-70; Bawden et al., ibid., 23, 1942, 314-328; Cohen, Proc. Soc. Exp. Biol, and Med., 4S, 1941, 163-167; Lea, Nature, U6, 1940, 137-138; Pirie et al., Parasitol., 30, 1938, 543-551; Price, Am. Jour. Bot., 25, 1938, 603-612; Am. Jour. Bot., 27, 1940, 530-541; Arch. Virusf., 1, 1940, 373-386; Price and Wyckoff, Phytopath., 29, 1939, 83-94; Smith, Parasitol., 29, 1937, 70-85; 29, 1937, 86-95; Smith and Bald, Parasitol., 27, 1935, 231-245; Smith and MacCle- ment, Parasitol., 32, 1940, 320-332. 5a. Marmor dodecahedron H. (loc.cit., 30). From Greek dodekahedron, dodeca- hedron. Common name : Tomato bushy-stunt virus. Hosts : SOLAN ACE AE — Lycopersicon esculentum Mill., tomato. Experiment- ally, also SOLA N ACE AE— Datura stra- monium L.; Nicotiana glutinosa L.; A. langsdorffii Weinm.; A^. tabacum L., to- bacco; Solarium nigrum L. LEGU- MINOSAE—Phaseolus vulgaris L., bean ; Vigna sinensis (L.) Endl., cowpea. COMPOSIT AE— Zinnia elegans .Jacq., zinnia. Geographical distribution : British Isles. 1170 MANUAL OF DETERMINATIVE BACTEEIOLOGY Induced disease : In tomato, some primary lesions necrotic, ring-like or spot-like, others masked, or disclosed only by chlorophyll retention in yellowing leaves. In young plants, systemic ne- crotic lesions may cause death ; in older plants, growth ceases, young leaves be- come pale j'ellow ; growing points may die, inducing growth of axillary buds to produce a bushy top ; older leaves become yellowed and show some purple colora- tion. In White Burley tobacco, local necrosis only, lesions small, red at first, then white. In cowpea, reddish necrotic primary lesions only. Transmission: By inoculation of e.\- pressed juice. Through dodder, Cuscvta campestris Yuncker {COA'VOLVULA- CEAE). Not through seeds of diseased plants. No insect vector is known. Serological relationships : A specific antiserum, prepared by a single intra- venous injection of rabbits with 2 mg of purified virus, gives granular, compact precipitates, serving for quantitative es- timation of this virus, antiserum being used at dilutions of 1 :200 or 1 :800, virus at dilutions to 10"". Immunological relationships: Will in- fect plants previously invaded by tobac- co-mosaic virus, tomato spotted-wilt virus, tobacco-ringspot virus, and Ber- gerac-ringspot virus. Filterability : Passes membranes down to 40 millimicrons average pore diameter. Other properties: Virus crystallizes from solutions of ammonium sulfate as isotropic, rhombic dodecahedra, which shrink and swell reversiblj' on drying and rewetling; shrinkage reduces size to 80 per cent of the wet dimensions. In the presence of heparin, non-birefringent prisms, rather than dodecahedra, appear. 820° = 132 X 10-13 cm. sec.-i dyne"!. Particle approximately spherical, 27.4 millimicrons in diameter by X-ray meas- urements (average diameter by filtration data, 14 to 20 millimicrons). Solutions do not show anisotropy of flow. Inac- tivated by drying. Molecular weight 8,800,000. Density 1.353. Molecular weight may be as high as 24,000,000 in solution, but the density is then lower, 1 .286. Analysis : carbon 47 to 50 per cent , nitrogen 15.8 to 16.4 per cent, phosphorus 1.3 to 1.5 per cent, ash 1.7 to 5 per cent, hj^drogen 7.2 to 8.2 per cent, sulfur 0.4 to 0.8 per cent, carbohydrate 5 to 6 per cent. Literature: Ainsworth, Jour. Ministry Agr., 43, 1936, 266-269 ; Bawden and Pirie, Nature, HI, 1938, 513; Brit. Jour. Exp. Path., 19, 1938, 251-263; Bernal and Fankuchen, Jour. Gen. Physiol., 25, 1941, 111-165; Bernal et al., Nature, lJt2, 1938, 1075; Cohen, Jour. Biol. Chem. lU, 1942, .353-362; Proc. Soc. Exp. Biol, and Med., 51, 1942, 104-105; Lauffer, Jour. Phys. Chem., U, 1940, 1137-1146; LaufTer and Stanley, Jour. Biol. Chem., 135, 1940, 463-472; Neurath and Cooper, Jour. Biol. Chem., 1S5, 1940, 455-462; Smith, Nature, 135, 1935, 908; Ann. Appl. Biol., 22, 1935, 731-741 ; Jour. Roy. Hort. Soc, 60, 1935, 448-451; Smith and MacClement, Para- sitol., 33, 1941, 320-330; Stanley, Jour. Biol. Chem., 135, 1940, 437-454. Key to the species of the Tobacco-Etch Virus Group. Viruses relatively susceptible to heat inactivation (inactivated at 52 to 58° C in 10 minutes). A small, closely allied group, tending to replace or to be replaced by each other, if present in mixture in tobacco. I. Not replaced, if in mixture, by other viruses of this group ; dominant member of the group in tobacco. 6. M armor erodens. II. Replaced by No. 6, not by No. 8, if in mixture with it in tobacco. 7. Marmor hyoscyami. III. Replaced by No. 6 or 7 if in mixture with either in tobacco. 8. Marmor upsilon. FAMILY MARMORACEAE 1171 6. Marmor erodens Holmes. (Holmes, Handb. Phytopath. Viruses, 1939, 40; Foliopellis erodens Valleau, Phytopath., 30, 1940, 825.) From Latin erodere, to erode or gnaw away. Common name: Tobacco-etch virus. Hosts : SOLAN ACE AE— Capsicum frutescens L., pepper ; Datura stramonium. L., Jimson weed ; Lycopersicon esculentum Mill., tomato; Nicotiana tabacum L., to- bacco; Petunia sp., petunia; Physalis heterophylla Xees. Geographical distribution: United States. Induced disease : In tobacco, systemic mild-mottling chlorosis, with traces of necrotic etching; intranuclear crystalline inclusions and intracytoplasmic granular and amorphous inclusions that tend to crystallize, forming needle-shaped bire- fringent bodies, 2 to 10 microns in length. Transmission : Experimentally, by My- zus persicae (Sulz.), M. circumfiexus (Buckt.), Aphis rhamni Boyer, A.fabae (Scop.), and Macrosiphum gei (Koch) (APHIDIDAE) ; by inoculation of ex- pressed juice. Serological relationships : Precipitin re- actions with homologous antisera, but no cross -react ions with tobacco-mosaic virus, tobacco -ringspot virus, potato - mottle virus, potato aucuba-mosaic virus, potato mild-mosaic virus, hs'oscyamus- mosaic virus, potato-veinbanding virus, or pea-mosaic virus. Immunological relationships : Protects tobacco against subsequent infection by potato-veinbanding virus and hyoscya- mus-mosaic virus. In mixed infections, 't suppresses and replaces these two viruses. Thermal inactivation : At 53 to 55° C in 10 minutes. Filterability : Passes Pasteur-Cham- be rland Li, not La, filter candle. Other properties : Sedimentation con- stant 820° = 170 X 10-13 cm. sec.-i dyne-i. Concentrated preparations show aniso- tropy of flow, indicating elongated parti- cle shape. Literature : Bawden and Kassanis, Ann. Appl. Biol., 28, 1941, 107-118; Fernow, Cornell Agr. Exp. Sta. (Ithaca), Mem. 96, 1925; Holmes, Phytopath., 32, 1942, 1058-1067; Johnson, Kentuckj^ Agr. Exp. Sta., Res. Bull. 306, 1930. Strains : A distinctive severe -symptom strain, i.solated from plants infected in nature and studied intensively, has been distinguished from the type, var. vulgare H. {loc. cit., 40), by the varietal name : 6a. Marmor erodens var. severum H. {loc. ci'L, 41 ) . Differing from the type by a tendency to induce more pronounced necrotic etching and a greater stunting effect in infected tobacco. 7. Manner hyoscyami spec. nov. From Xew Latin Hyoscyamus, genus name of plant from which this virus was first isolated. Common names : Hyoscyamus-mosaic virus, Hy. Ill virus, Hyoscyamus-III- disease virus. Hosts : SOLAN AC EAE— Hyoscyamus niger L., henbane. Experimentally, also Nicotiana tabacum L., tobacco. Insusceptible species -.CUCURBITA- CEAE — Cucumis sativus L., cucumber. Geographical distribution: England. Induced disease : In henbane, chlorotic clearing of veins followed by yellow-mot- tling mosaic. Transmission: By inoculation of ex- pressed juice to dilutions of 10^^. B3- aphids, Myzus persicae (Sulz.), M. cir- cumfiexus (Buckt.), and Macrosiphum solanifolii Ashm. (= M. gei Koch) (APHIDIDAE). Serological relationships : Several iso- lates of this virus give mutual cross- precipitin reactions but no precipitation occurs when antiserum prepared with this virus is mixed with cucumber-mosaic virus, tobacco-etch virus, or potato-vein- banding virus. Immunological relationships : Xo im- munity with respect to this virus is in- duced in tobacco by previous infection with cucumber-mosaic virus. Potato- veinbanding virus is unable to multiply in the presence of this virus and is re- placed by it. Tobacco-etch virus pro- 1172 MANUAL OF DETERMINATIVE BACTERIOLOGY tects against this virus and replaces it in mixed infections. Thermal inactivatibn : At 58° C in 10 minutes. Filterability : Passes Chamberland Li, but not Ls, filter candles. Other properties : Concentrated solu- tions show anisotropy of flow. Yield of virus, 1 to 3 mg per liter of juice expressed from diseased tobacco plants. Literature : Bawden and Kassanis, Ann. Appl. Biol., 28, 1941, 107-118; Hamilton, ibid., 19, 1932, 550-567; Sheffield, ibid., 25, 1938, 781-789; Watson and Roberts, Proc. Roy. Soc. London, Ser. B, 127, 1939, 543-576. 8. Marmot upsilon comb. nov. {Mar- mor cucumeris var. vpsilon Holmes, loc. cit., 33; Murialba vcnataenia Valleau, Phytopatli., 30, 1940, 824.) From Greek name of the letter Y, sometimes used to denote this virus. Common names : Potato -veinbanding virus, potato virus Y. Hosts: SOLAN ACE AE—Solaniim tu- herosxim L., potato; Nicotiana tabacum L., tobacco. Experimentally, also Ltj- cium barbariim L. Geographical distribution: England, France, United States, Brazil. Rare in Scotland and part of Ireland. Induced disease : In some potato varie- ties, leaf drop and necrotic stem-streak; in others, no signs of disease; in still others, chlorotic mottling with or without necrosis. In combination with strains of the ]3otato-mottle virus {Marmor dubium), this virus causes rugose mosaic, a common and destructive double-virus disease . Transmission: By inoculation of ex- pressed juice. By aphid, Myzus persicae (Sulz.); experimentally, also by Aphis rhamni Boyer (synonym for Aphis ab- breviata Fatch) (APHIDIDAE). Serological relationships : Precipitin re- actions with homologous antisera. No cross reactions with tobacco-mosaic virus, tobacco-etch virus, hyoscyamus -mosaic virus, potato-mottle virus, potato mild- mosaic virus, potato aucuba-mosaic virus, tobacco-ringspot virus, or common pea- mosaic virus. Reported cross reaction with cucumber-mosaic virus needs con- firmation. Immunological relationships : A mild strain protects against subsequent infec- tion with the typical virus. This virus is suppressed and replaced by hyoscya- mus-mosaic virus and by tobacco-etch virus in mixed infections. Thermal inactivation : At 52° C in 10 minutes. Filterability : Passes with difficulty through Gradocol membrane of 42 milli- micron average pore diameter. Other properties : Inactivated by dry- ing. Literature: Dennis, Nature, 14-2, 1938, 154; Johnson, Phytopath., 25, 1935, 650-652; Jones and Vincent, Jour. Agr. Res., 55, 1937, 69-79; Kassanis, Ann. Appl. Biol., 29, 1942, 95; Koch, Phyto- path., 28, 1933, 319-342; Kramer and Sil- berschmidt, Arquivos Inst. Biol., Sao Paulo, Brazil, 11, 1940, 165-188; Salaman, Nature, 139, 1937, 924; Smith, Proc. Roy. Soc, Ser. B, 109, 1931, 251-267; Smith and Dennis, Ann. Appl. Biol., 27, 1940, 65-70. Key to the species of the Cuciunber -Mosaic Virus Group. Viruses relatively susceptible to heat inactivation, requiring less than 10 minutes at 85 to 90° C for complete inactivation. Not replacing potato-veinbanding virus in mixed infections. I. Infecting both dicotyledonous and monocotyledonous plants. 9. Marmor cucumeris. II. Infecting dicotyledonous, but not monocotyledonous, plants. 10. Marmor solani. 11. Marmor aucub a. 12. Marmor umbelliferarum . FAMILY MARMORACEAE 1173 13. M armor cruciferarum. 14. M armor brassicac. 15. Marmor betae. 16. Marmor lactucae. 17. Marmor dahliae. 18. Marmor phaseoli. 19. Marmor leguminosarum. 20. Marmor pisi. 21. Marmor medicaginis. III. Infecting monocotyledonous, but not dicotyledonous, plants. 22. Marmor tulipae. 23. Marmor mite. 24. Marmor iridis. 25. Marmor sacchari. . 26. Marmor cepae. 27. Marmor scillearum. 9. Marmor cucumeris Holmes. (Holmes, Handb. Phytopath. Viruses, 1939, 31; Murialba cucumeris Valleau, Phytopath., 30, 1940, 823.) From Latin ciicumis, cucumber. Common name : Cucumber-mosaic virus. Hosts : Very wide range of hosts among dicotyledonous and monocotyledonous plants; cucumber, celery, spinach, to- bacco, and pepper are sometimes serioush' affected. Overwintering hosts are : SOL- ANACEAE — Physalis subglabrata Mac- kenzie and Bush, P. helerophylla Nees. ASCLEPIADACEAE—Asclepias syri- aca L. PHYTOLACCACEAE— Phyto- lacca decandra L. LAB I AT AE — Ne- peta cataria L. Probably there are also other susceptible perennials. Geographical distribution : Probably al- most world-wide. Induced disease : In cucumber, Cucu- mis sativus L., yellowish-green systemic mottling. Leaves small, distorted, curled; plants dwarfed, internodes short- ened. Few fruits set. Fruits mottled, nusshapen, giving the disease the name "white pickle." In black cowpea, Vigna sinensis (L.) Endl., small reddish ne- crotic local lesions only. No intracellular bodies are found in plants infected with cucumber-mosaic virus. Transmission : By inoculation of ex- pressed juice. By aphids, Myzus persi- cae (Sulz.), M. pseudosolani Theob., M. circumflexus (Buckt.), Macrosiphum sol- anifolii Ashm., and Aphis gossrjpii Glov. {APHIDIDAE). Through seeds of dis- eased plants in Echinocystis lohata (Michx.) Torr. and Gray, wild cucumber, in Ciicumis melo L., muskmelon, and in Cucurbita pepo L., vegetable maiTow. By several species of dodder, Cusciita calif ornica Choisy, C. campeslris Yuncker, and C. suhinclusa Dur. and Hilg. {CONVOLVULACEAE). Immunological relationships : Infection with the type and other chlorotic -mot- tling strains protects zinnia against sub- sequent infection by an indicator strain of this virus (var. judicis). Thermal inactivation : At 70 to 80'^ C in 10 minutes. Filterability : Passes Berkefeld W and N filters and collodion membranes of 45 millimicron average pore diameter. Other properties : Inactivated by dry- ing or 3 to 4 days' storage in juice at room temperature. Literature: Ainsworth, Ann. Appl. Biol., 25, 1938, 867-869; Chamberlain, New Zealand Jour. Science and Tech- nology, 21, 1939, 73A-90A ; Celino, Philip- pine Agr., 29, 1940, 379-414; Doolittle, Phytopath., 6, 1916, 145-147; U. S. Dept. Agr., Bull. 879, 1920; Doolittle and Walker, Jour. Agr. Res., 31, 1925, 1-58; Gilbert, Phytopath., 6, 1916, 143-144; Hoggan, Jour. Agr. Res., 47, 1933, 689- 704; Jagger, Phytopath., 6, 1916, 148-151 ; 1174 MANUAL OF DETERMINATIVE BACTERIOLOGY 8, 1918, 32-33; Kendrick, Phytopath., 24, 1934, 820-823; Mahoney, Proc. Am. Soc. Hort. Sci., 332, 1935, 477-480; Price, Phytopath., 25, 1935, 776-789; 29, 1939, 903-905; Am. Jour. Bot., 27, 1940, 530- 541; Storey, Ann. Appl. Biol., 26, 1939, 298-308. Strains : Various host plants seem to have induced specialization of cucumber- mosaic virus in strains particularly adapted to existence in their tissues. Several of these and certain laboratory- derived strains useful in technical proce- dures have been distinguished from the type, var. imlgare H. {loc. cit., 31), by varietal names, as follows : 9a. Marnior cucumeris var. commelinac H. {loc. cit., 35). From New Latin Commelina, generic name of weed serving as a natural reservoir of this strain. Common name : Southern celery -mosaic strain of cucumber-mosaic virus. Differ- ing from the type in severity of disease induced in celery and some other plants. Transmitted by Aphis gossypii Glov., A . maidis Fitch, and Pentalonia nigro- nervosa Coq. (APH IDIDAE) . (Price, Phytopath., 25, 1935, 947-954; Wellman, ibid., 24, 1934, 695-725, 10.32-10.37; 25, 1935, 289-308, 377-404.) 9b. Marmor cucumeris var. phascoli H. (loc. cit., 36). From New Latin Pha- seolus, generic designation of lima bean. Common name: Lima-bean strain of cu- cumber-mosaic virus. Differing from type of species in ability to cause a chlo- rotic mottling disease in lima bean in nature. (Harter, Phytopath., 26, 1936, 94; Jour. Agr. Res., 56, 1938, 895-906; McClintock, Phytopath., 7, 1917, 60.) 9c. Marmo7- cucumeris var . lilii YL. {loc. cit. ,37). From Latin /(7i'H?«, lily. Com- mon name : Lily-mosaic strain of cucum- ber-mosaic virus. Differing from the type variety by ability to persist in na- ture in lilies, producing masked infection or chlorotic mottling unless in mixture with lily-symptomless virus {Adelonosus lilii), when a more severe disease involv- ing necrosis is induced. (Brierley, Phy- topath., 29, 1939, 3; 30, 1940, 250-257; Brierley and Doolittle, ibid., 30, 1940, 171-174; Ogilvie and Guterman, ibid., 19, 1929, 311-315; Price, ibid., 27, 1937, 561-569.) 9d. Marmor cucumeris var. judicis H. {loc. cit., 38). From Latin jwdex, judge. Common name : Indicator strain of cu- cumber-mosaic virus. Differing from the type variety in inducing the forma- tion of necrotic local lesions in zinnia {Zinnia elegans Jacq., COMPOSITAE). Previous infection of zinnia by other strains of cucumber-mosaic virus in- hibits the formation of these necrotic local lesions, identifying the strains as related to each other and to the indicator strain. (Price, Phytopath., 24, 1934, 743-761 ; 25, 1935, 776-789.) 9e. Marmor cucumeris var. vignae H. {loc. cit., 39). From New Latin Vigna, generic name of cowpea. Common name : Cowpea-mottling strain of cucumber- mosaic virus. Differing from the type variety in producing systemic chlorotic mottling, rather than reddish-brown ne- crotic local lesions, in Black cowpea. Not known in nature but derived experi- mentally from a mild-mottling strain of cucumber-mosaic virus during serial passage in cowpea. (Price, Phytopath., 24, 1934, 743-761 ; 25, 1935, 776-789.) 10. Marmor solani H. {loc. cit., 47). From New Latin Solanum, generic name of potato. Common names : Potato mild-mosaic virus, potato virus A. Hosts: SOLAN AC EAE— Solanum. tu- berosum. L., potato. Experimentally, also Nicotiana tabacum L., tobacco; Sol- anum nigrum L. var. nodiflorum; and Datura stramonium L., Jimson weed. Geographical distribution : United States, England, Holland. Induced disease : In potato, very mild chlorotic mottling or masked symptoms in some varieties (as Irish Chieftain), systemic necrosis in others (for example," FAMILY MARMORACEAE 1175 British Queen). Immunity to aphid in- fection with this virus is found in the varieties Katahdin and Earlaine. A combination disease, characterized by pronounced yellow-mosaic patterns, is caused by this virus in the variety Irish Chieftain if the potato-veinbanding virus {Marrnor wpsilon) is also present. In tobacco, experimentally, faint veinband- ing mosaic. Transmission: To potato, by rubbing methods of inoculation of expressed juice, using carborundum powder; to tobacco, by rubbing without carborundum. By aphids. Aphis abbreviata Patch and Myzus persicae (Sulz.) (APHIDIDAE). Serological relationships : No cross - precipitin reactions with potato aucuba- mosaic virus, potato-veinbanding virus, tobacco-mosaic virus, tobacco-etch virus, tobacco-ringspot virus, or pea-mosaic virus . Immunological relationships : A feeble strain of this virus has been found to pro- tect fully against the typical strain in the Netherlands. Thermal inactivation : At 50° C in 10 minutes . Literature: Bawden, Ann. Appl. Biol., ^3, 1936, 487-497; Chester, Phytopath., 25, 1935, 686-701; Dykstra, Phytopath., 29, 1939, 40-67; Hansen, Tidsskr. Plan- teavl, 42, 1937, 631-681; Murphy and Loughnane, Sci. Proc. Roy. Dublin Soc, 21, 1936, 419-430; Murphy and McKay, ibid., 20, 1932, 227-247; Oortwijn Botjes, Tijdsch. Plantenziekten, 45, 1939, 25- 29; Schultz ei al., Phytopath., 27, 1937, 190-197; SO, 1940,944-951. 11. Maimer aucuba H. {loc. cit., 49). From New Latin Aucuba, a genus of plants having mottled foliage . Common name : Potato aucuba -mosaic virus. Hosts: SOLAN ACE AE—Solanum tu- berosum L., potato. Experimentally, also Atropa belladonna L. (symptomless) ; Capsicum frutescens L., pepper; Datura stramonium L., Jimson weed (symptom- less); Hyoscyamus niger L., henbane (symptomless); Lycopersicon esculentum Mill., tomato; Petunia hybrida Vilm., petunia (symptomless); Nicotiana ta- bacum L., tobacco (symptomless) ; *Soia- num dulcamara L., bittersweet ; S. nigrum L. var. nodiflorum. Geographical distribution : United States, Great Britain, Europe. Induced disease : In potato, yellow spots on lower leaves of some varieties ; in the variety Irish Chieftain, brilliant yellow mottle over whole plant, perhaps because of simultaneous presence of po- tato mild-mosaic virus in this variety. Necrosis of the cortex and of the pith in tubers in many varieties. Transmission : By inoculation of ex- pressed juice. Probably by aphid, My- zus persicae (Sulz.) {APHIDIDAE). Serological relationships : No precipitin cross -reactions with potato mild-mosaic virus, potato-veinbanding virus, tobacco- mosaic virus, tobacco-etch virus, tobacco- ringspot virus, or pea-mosaic virus. Precipitin cross-reactions with the Canada-streak strain of potato aucuba- mosaic virus. Thermal inactivation : At 65 to 68° C in 10 minutes. Filterability : Passes Pasteur-Cham- berland Li filter, but not L3 or L5. Literature: Chester, Phytopath., 25, 1935, 686-701; 27, 1937, 903-912; Clinch, Sci. Proc. Roy. Dublin Soc, 22, 1941, 435-445; Clinch et al., ibid., 21, 1936, 431- 448; Dykstra, Phytopath., 29, 1939, 917- 933. Strains : One strain differing from the t3'pe has been given a varietal name : 11a. Marmor aucuba var. canadense Black and Price. (Phytopath. 30, 1940, 444.) From common name of strain. Common name : Canada-streak strain of potato aucuba-mosaic virus. Differing from the type variety by tendency to produce necrosis in stems, veins, petioles, and leaves and also, about 2 months after harvest, in pith of tuber, especially at stem end. (Chester, Phytopath., 27, 1176 MANUAL OF DETERMINATIVE BACTERIOLOGY 1937, 903-912; Dykstra, Phytopath., 29, 1939, 917-933.) 12. Marmor umbelliferarum H. {loc. cit.,&7). From New Latin Umbelliferae, family name of plants among which celery is classified. Common name : Celery -mosaic ^irus, western celery -mosaic virus. Hosts : UMBELLIFERAE— A-pium graveolens L., celery and celeriac ; Daucns carota L., carrot. Experimentally, also Aneihum graveolens L., dill; Anthriscxm cerefolium (L.) Hoffm., salad chervil; Carum carvi L., caraway; Coriandrum sativvm L., coriander; Pitroselinnm hor- tense Hoffm., parsley. Insusceptible species: Cucumis satimts L., cucumber, and all other tested species not of the family Umbelliferae. Geographical distribution: United States (California). Induced disease : In celery, at first, clearing of veins in young leaves; later, foliage yellowed, plmt stunted, young petioles shortened, older petioles hori- zontal, giving plant a flat appearance. Foliage mottled green and yellow ; leaflets narrow, twisted or cupped; older leaves with some necrosis; petioles with white streaks or spots. In celeriac, clearing of veins, followed by systemic chlorotic mottling. In carrot, chlorotic spotting of young leaves, followed by systemic chlorotic mottling. Transmission : Bj' inoculation of ex- pressed juice, in dilutions to 1:4000. No specific insect vector is known, but 11 species of aphids capable of breeding on celery transmit the virus, though they do not long retain the power of transmission after leaving diseased plants. These vec- tors are Aphis apigraveolens Essig, A. apii Theob., A. fcrriiginea-striata Essig, A. (jossypii Glov., A. rniddlcionii Tho- mas, A. ramicis Linn., CavarieUa cap- reae (Fabr.), Myzufi circnynflcxu-'^ (Buckt.), .1/. convolvuli (Kalt.), M. per- sicae (Sulz.), Rhopalosiphvm mclliferuin (Hottes) (APHIDIDAE). Some aphids not able to breed on celery also transmit this virus. Thermal inactivation : At 55 to 60° C in 10 minutes. Filterability : Passes all grades of Chamberland filters. Other properties : Virus active after storage at —18° C for 18 months. Literature : Severin and Freitag, Hil- gardia,^^, 1938, 493-558. 13. Marmor cruciferarum H. (loc. oil., 69). From New Latin Cruciferae, family name of plants among which cauliflower is classified. Common name : Cauliflower-mosaic virus. Hosts: CRUCIFERAE — Brassica ol- eracea L., cauliflower, kale, Brussels sprouts, cabbage, and broccoli; B. cam- pestris L., wild yellow mustard; Mat- thiola incana R. Br., annual stock. Ex- perimentally, also Brassica adpressa Boiss; B. alba Rabenh., white mustard; B. arvensis (L.) Ktze., charlock; B. juncea Coss., leaf mustard (one strain not susceptible); B. napus L., rape; B. pe- isai Bailey, pe-tsai ; B. nigra Koch, black mustard; B. rapa L., turnip; Capsella bursa-pastoris Medic, shepherd's purse; Iberis amara L., rocket candytuft; Lepi- dium sativum L., garden cress; Lxinaria annua L., honesty; Raphanns raphanis- trum L., white charlock; R. saiivvs L., radish. Insusceptible species : CHENOPODI- ACE AE—Spinacia oleracea L. COM- POS IT AE—Lactuca saliva L. CR UCI- FERAE — Alyssinn saxatile L.; A. maritimum Lam.; Arabis albida Stev.; Athysanus pusillus Greene; Brassica juncea Coss. (Japanese strain; another strain susceptible); Cheiranthus cheiri L.; Erysivnivi perojsldanum Fisch. and Mey. ; Hespcris malronalis L. ; Malcomia mariiima R. Br. ; Roripa nasturtium liushy ; Stanleya pinnata (Pursh.) Britt.; Thysanocarpus radians Benth. LEGU- MINOSAE—Vicia faba L. SOLAN A- CEAE — Capsicum Jrutcsccns L. ; Lycoper- FAMILY M ARMOR ACEAE 1177 sicon esculenlum Mill.; L. pimpinellifol- ium Mill.; Nicotiana glutinosa L.; A'. langsdorffii Weinm. ; N. tabacum L. vars. Turkish and White Burley. TROPAE- OLACEAE — Tropaeoliun majus L. UMBELLIFERAE—Apium graveo- lens L. Geographical distribution : United States, England. Induced disease : In cauliflower, clear- ing of veins, followed by mild chlorotic mottling, veins usually banded with, dark green, necrotic flecks later in chlorotic areas. Midrib curved, leaves distorted. Plant stunted; terminal head or curd dwarfed. Solanaceous plants appear to be immune, a point of distinction between this virus and turnip-mosaic virus, Mai'- mor brassicae. Transmission : By inoculation of ex- pressed juice, using carborundum pow- der. By many aphid species, Brevicorync brassicae (Linn.), cabbage aphid; Rhopa- losiphum pseudobrassicae Davis, false cabbage aphid; Myzus persicae (Sulz.), peach aphid; Aphis graveolens Essig, celery leaf aphid; ^4. apigraveolens Essig, celery aphid; A. middletonii Thomas, erigeron root aphid; A. gossypii Glov., cotton aphid; Cavariella capreae (Fabr.), yellow willow aphid; Myzus circumflexus (Buckt.), lily aphid; Rhopalosiphum mel- liferum (Hottes), honeysuckle aphid. (APHIDIDAE). No seed transmission. Thermal inactivation : At 75^ C in 10 minutes. Literature: Caldwell and Prentice, Ann. Appl. Biol., 29, 1942, 366-373, 374- 379; Rawlins and Tompkins, Phytopath., U, 1934, 1147 (Abst.); Tompkins, Jour. Agr. Res., 55, 1937, 33-46. 14. Manner brassicae H. (H., loc. cit., 70; Martnor matthiolae H., loc. cit., 71.) From New Latin, Brassica, generic name of turnip. Ccmmon name: Turnip-mosaic virus. Hosts : CR UC IF ERAE— Brassica rapa L., turnip; B. napobrassica Mill., swede or rutabaga; B. napus L., rape; B. nigra (L.) Koch, black mustard; B. oler- acea L., cabbage; Armoracia rusticana Gaertn., horse-radish; Cheiranthus cheiri L., w'allflower; Maithiola incana R. Br., stock; Sinapis alba L., white mustard. Experimentally, also CRUCIFERAE— Berteroa incana (L.) DC.; Brassica alba Rabenh., white mustard; B. arvensis (L.) Ktze. ; B. chinensis L., Chinese cab- bage; B. juncea (L.) Coss.; Capsella bursa-pastoris (L.) Medic; Cardamine heterophylla (Forst. f.) O. E. Schultz; Cheiranthus allionii Hort.; Coronopus didymus Smith; Hesperis matronalis L.; Lepidium ruderale L.; L. sativum L., L. virginicum L.; A^asturtium officinale R. Br.; N eslia paniculata (L.) Desv.; Radi- cula palustris (L.) Moench.; Raphanus sativus L.; Sisymhrium altissimum L.; S. officinale (L.) Scop.; Thlaspi arvense L. CHENOPODIACEAE—Beta vul- garis L.; Spinacia oleracea L., spinach. COM POS I TAE— Calendula officinalis L. Zinnia elegans Jacq. RANUNCULA- CEAE — Delphinium ajacisL. SOLAN A- CEAE — Lycopersicon pimpinellifolium Mill.; Nicotiana bigelovii S. Wats.; A^. glutinosa L.; A', langsdorffii Weinm.; A''. repanda Willd. ; N. rustica L. ; A', sylves- tris Speg. and Comes; A", tabacum L., tobacco; Petunia hybrida Vilm. Geographical distribution ; United States, England, New Zealand. Induced disease : In turnip, systemic chlorotic mottling; plants stunted, leaves distorted. In tobacco, experimentally, characteristic necrotic primary lesions only. Transmission : Bj^ inoculation of ex- pressed juice. By cabbage aphid, Brevi- corync brassicae (Linn.), and by the peach aphid, Myzus persicae (Sulz.) (APHIDIDAE). Thermal inactivation : At 54° C in 10 minutes . Strains : A considerable number of strains of this virus appear to occur in nature, but those that have been studied often have been considered as distinct viruses and not compared with each other 1178 MANUAL OF DETERMINATIVE BACTERIOLOGY under identical circumstances. More work is needed to show existing alliances. Literature : Chamberlain, New Zealand Jour. Agr., 53,1936, 321-330; New Zealand Jour. Science and Technology, 21, 1939, 212A-223A; Clayton, Jour. Agr. Res., 40, 1930, 263-270; Gardner and Kendrick, ibid., 22, 1921, 123-124 ; Hoggan and John- son, Phytopath., 25, 1935, 640-644 ; Larson and Walker, Jour. Agr. Res., 59, 1939, 367-392; 62, 1941, 475-491 ; Schultz, Jour. Agr. Res., 22, 1921, 173-178; Smith, Ann. Appl. Biol., 22, 1935, 239-242; Tompkins, Jour. Agr. Res., 57, 1938, 589-602; 58, 1939, 63-77; Tompkins et al., ibid., 57, 1938, 929-943. 15. Marmor betae H. {loc. cit., 72). From Latin beta, beet. Common name : Sugar-beet mosaic virus. Hosts : C HENOPODI ACE AE— Beta vulgaris L., beet; Spinacia oleracea L., spinach. Geographical distribution: France, Denmark, Germany, Sweden, L^nited States, England. Induced disease : In beet, discrete yellowish secondary lesions or clearing of veins on young leaves, followed by chlo- rotic mottling of newly formed leaves. Darkening of vascular tissue. Leaves bend back near tips, which sometimes die. Intracellular bodies formed. In spinach, 6 to 21 days after infection, chlorotic flecks on young leaves. Plant stunted, outer leaves killed, dying from their tips back. Center of plant survives for a time, but finally dies. Transmission: By inoculation of ex- pressed juice, in dilutions to 10~^ By aphids, Myzus persicae (Sulz.), Aphis rumicis Linn., and perhaps Macrosiphuni solanifolii Ashm. (= M. gei Koch) (APHIDIDAE). No seed transmis- sion. Thermal inactivalion : At 55 to 60° C in 10 minutes. Other properties : Inactivated bj^ standing in expressed juice for 24 to 48 hours at about 70° F. Literature : Boning, Forsch. Geb. Pflan- zenkr. u. Immun. Pflanzenreich, 3, 1927, 81-128; Cent. f. Bakt., II Abt., 71, 1927, 490-497; Gratia and Manil, Compt. rend. Soc. Biol., Paris, 118, 1935, 379-381; Hoggan, Phytopath., 23, 1933, 446-474; Jones, Washington Agr. Exp. Sta. Bull. 250, 1931; Lind, Tidsskr. Planteavl, 22, 1915, 444-457; Robbins, Phytopath., 11, 1921, 349-365; Schmidt, Ber. Deutsch. Bot.Ges., 45, 1927,598-601. 16. Marmor lactucae H. {loc. cit., 84). From Latin lactuca, lettuce. Common name : Lettuce-mosaic virus. Hosts: COM POS I TAE— Lactuca sa- liva L., lettuce; Senecio vulgaris L., groundsel. Experimentally, also COM- POSITAE—Sonchus asper Hoffm., prickly sow-thistle. LEGUMINOSAE — Lathyrus odoratus L., sweet pea; Pisum sativinn L., pea. Insusceptible species : COMPOSITAE — Sonchus oleraceus L., S. arvensis L., Taraxacum officinale Web., Carduus ar- vensis Curt. CRIJCIFERAE—Bras- sica oleracea L. CUCURBIT ACE AE— Cucumis sativus L. SOLAN ACE AE — Lycopersicon esculentum Mill., Nicotiana tahacum L., N. glutinosa L., Datura stramonium L. Geographical distribution : United States, England, Germany, Bermuda. Induced disease: In lettuce varieties, clearing of veins followed by systemic chlorotic mottling, dwarfing and defective hearting; sometimes by scorching of leaf edges, vein necrosis or necrotic flecking between veins. Transmission : By inoculation of ex- pressed juice, in dilutions to 1:100 if mixed with a little 0.5 per cent sodium sulphite solution and a trace of powdered carborundum. By aphids, Myzus persi- cae (Sulz.) and Macrosiphum gei Koch (APHIDIDAE). Through seeds from diseased plants. It is believed that seed- borne virus is the most important source of primary inoculum in the spring. Thermal inactivation : At 55 to 60° C in 10 minutes. FAMILY ]VL\RMORACEAE 1179 Filterability : Fails to pass Li Pasteur- Chamberland filter. Literature: Ainsworth and Ogilvie, Ann. Appl. Biol., 26, 1939, 279-297; Jag- ger, Jour. Agr. Res., 20, 1921, 737-740; Newhall, Phytopath., 13, 1923, 104-106. 17. Marmor dahliae H. (loc. cit., 85). From Xew Latin Dahlia, generic name of host plant. Common name : Dahlia-mosaic virus. Hosts: COMPOSITAE— Dahlia pin- nata Cav., dahlia. Experimentally, also D. imperialis Roezl. ; D. maxonii Safford. Geographical distribution: L'nited States, Holland, Germany, England. Induced disease : In intolerant varieties of dahlia, chlorotic mottling of foliage, leaf distortion, dwarfing of all stems and of roots, occasionalh^ necrotic streaking of midveins. In tolerant varieties, incon- spicuous chlorotic mottling or masked symptoms . Transmission : Bj- aphid, Myzus persi- cae (Sulz.) (APHIDIDAE). By graft- ing. Not by inoculation of expressed juice. Not through soil. Not through seeds from diseased plants. Literature: Brierley, Am. Dahlia Soc. Bull., Ser. 9, No. 65, 1933; Contrib. Boyce Thompson Inst., 5, 1933, 23.5-288; Gold- stein, Bull. Torrey Bot. Club, 54, 1927, 285-293. 18. Marmor phaseoli H. {loc. cit., 87). From New Latin Phaseolus, generic name of bean. Common name : Bean-mosaic virus. Hosts : LEGUM I NOSAE— Phaseolus vulgaris L., bean. Experimentally, also Phaseolus acutifolius Gray var. latifolius Freem.; P. aureus Roxb.; P. calcaratus Roxb.; P. lunatus L.; Lespedeza striata (Thunb.) Hook, and Arn. ; Viciafaba L. ; V. sativa L., spring vetch. Insusceptible species: LEGUM I NO- SAE— Pisuni sativum L., garden pea; Lathyrus odoratus L., sweet pea. Geographical distribution : World-wide, wherever beans are grown . Induced disease : In bean, first leaves to be affected are crinkled, stiff, chloro- tic; later leaves show chlorotic mottling; leaf margins often rolled down. Opti- mum temperature for expression of dis- ease, 20 to 28° C, partial masking at 28 to 32° C, complete masking at 12 to 18° C. Transmission : By inoculation of ex- pressed juice in dilutions to 1 :1000, using carborundum or other abrasive powder. By aphids, Aphis rumicis Linn., Macro- siphum (= Illinoia) solanifolii Ashm., M. pisi Kalt., Aphis gossypii Glov., A. medicaginis Koch, A. spiraecola, Brevi- coryne brassicae (Linn.), Hyalopterus atriplicis Linn., Macrosiphum ambrosias Thos., Rhopalosiphum pseudobrassicae Davis, and Myzus persicae (Sulz.) (APHIDIDAE). In beans, there is seed transmission to 30 to 50 per cent of plants grown from infected parents ; pollen from infected plants is said to transmit virus. Thermal inactivation : At 56 to 58° C in 10 minutes. Literature: Fajardo, Phytopath., 20, 1930, 469-494, 883-888; Murphy, ibid., SO, 1940, 779-784; Murphy and Pierce, ibid., 28, 1938, 270-273; Parker, Jour. Agr. Res., 52, 1936, 895-915; Pierce, Phy- topath., 24, 1934, 87-115; , Jour. Agr. Res., 49, 1934, 183-188; 51, 1935, 1017-1039; Reddick, II Congr. Intern. Path. Comp., 1931, 363-366; Reddick and Stewart, Phytopath., 8, 1918, 530-534; Richards and Burkholder, Phytopath., 33, 1943, 1215-1216; Wade and Andrus, Jour. Agr. Res., 63, 1941, 389-393; Wade and Zau- meyer, U. S. Dept. Agr., Circ. 500, 1938; Walker and Jolivette, Phytopath., 33, 1943, 778-788; Zaumeyer and Kearns, ibid., 26, 1936, 614-629; Zaumeyer and Wade, Jour. Agr. Res., 51, 1935, 715-749. 19. Marmor leguminosarum H. {loc. cit., 89). From New Latin Leguminosae , family name of pea. Common name: Pea-mosaic virus. Hosts : LEG UM I NOSAE— Lathyrus odoratus L., sweet pea; Pisum. sativum L., pea ; Trifolium pratense L., red clover ; Viciafaba L., broad bean. Experiment- 1180 MANUAL OF DETERMINATIVE BACTERIOLOGY ally, also Cicer arietinum L.; Desmodium canadense (L.) DC; Lathyrus sativus L., grass pea; Litpiwws alhus L., white lupine ; L. angiistijolius , blue lupine; L. densi- floriis Benth. ; L. hartwegii Lindl. ; L. na- nus Dougl.; Medicago arahica Huds., spotted bur clover; M. hispida Gaertn., toothed bur clover; Melilotus alba Desr., white sweet clover; M. indicaAU., annual yellow sweet clover; M. officinalis (L.) Lam., yellow sweet clover; Phaseolus acutijolius Gray, tepary bean; P. vulgaris L., bean; Trifolium agrarium L. ; T. caro- linianum Michx.; T. duhi^im Sibth.; 7'. glomeratum L., cluster clover; T. hy- bridurn L., alsike clover; T. incarnatinn L., crimson clover; T. procumbcns L. ; T. reflexum L.; T. suaveolens, Persian clover ; Vicia saliva L., common vetch. Insusceptible species: All tested spe- cies in families other than the Leguni- inosae. Geographical distribution: United States, British Isles, Europe, New Zealand . Induced disease : In pea, clearing of veins in young leaves, followed by chloro- sis of newly formed leaves, stunting of plant, and systemic chlorotic mottling. In sweet pea, systemic chlorosis and chlorotic mottling, flower colors broken. In lupine, necrotic streak on one side of stem, stunting of plant and bending of growing point to injured side. Plant soon wilts and dies. In Vicia faba, mot- tled leaves contain characteristic iso- metric crj^stals in host -cell nuclei (espe- cially within nucleoli) as well as in cell cytoplasm. • Transmission : By inoculation of ex- pressed juice, with ease. Bj' aphids, Macrosiphian pisi Kalt., M. solanifolii Ashm. (= M. gci Koch), and Aphis rumicis Linn. (APHIDIDAE). Not transmitted through seed. Serological relationships : Specific pre- cipitin reactions differentiate this virus from tobacco -mosaic virus, tobacco-etch virus, potato-mottle virus, potato mild- mosaic virus, potato aucuba-mosaic virus, and tobacco -ringspot virus. Thermal inactivation : At 60° C in 10 minutes. Literature: Chester, Phytopath., 25, 1935, 68G-701 ; Doolittle and Jones, ibid., 15, 1925, 763-772 ; Johnson and Jones, Jour. Agr. Res., 54, 1937, 629-638; McWhorter, Phytopath., 31, 1941, 760-761; Murphy and Pierce, ibid., 27, 1937, 710-721; Os- born, ibid., 27, 1937, 589-603; Pierce, Jour. Agr. Res., 51, 1935, 1017-1039; Spierenburg, Tijdschr. Plantenz., 4^, 1936, 71-76; Zaumeyer and Wade, Jour. Agr. Res., 53, 1936, 161-185. 20. Marmor pisi H. {loc. cit., 90). From Ivatin pisum, pea. Common name : Pea enation-mosaic virus. Hosts: LEGUMINOSAE— Pisum sa- tivum L., pea; Vicia faba L., broad bean. Experimentally, also Lathyrus odoratus L., sweet pea; Soja max (L.) Piper, soy bean; Trifolium incarnatum L., crimson clover. Insusceptible species: LEGUMINO- SAE— Arachis hypogaea L., peanut; Medicago sativa L., alfalfa; Melilotus alba Desr., white sweet clover; M. officinalis (L.) Lam., yellow sweet clover; Phaseo- lus aureus Roxb., mung bean; P. vulgaris L., bean; Trifolium hybridum L., alsike clover; T. pratense L., red clover; T. repens L., white Dutch clover. SOL- A NACEAE — Lycopersicon esculentum Mill., tomato; Solanum tuberosum L., potato . Geographical distribution: United States, perhaps Germany. Induced disease: In peas, systemic chlorotic mottling; in some varieties, as Alderman, occasional necrotic spots and numerous enations on lower surfaces of leaves. Pods distorted. In broad bean, systemic chlorotic spotting and striping of leaves. In sweet pea and soy bean, experimentally, systemic chlorotic mot- tling. Transmission : By inoculation of ex- pressed juice, using carborundum; more readily from aphid-inoculated plants than from mechanically -inoculated plants. FAMILY MARMORACEAE 1181 Infective in dilutions to 10~'. By aphids, Macrosiphum pisi Kalt. and M. solani- folii Ashm. (= M. gei Koch) {APHI- DIDAE), with incubation periods of about 12 hours before the insects can infect. Xot through seeds from diseased plants. Thermal inactivation : At 66° C in 10 minutes. Literature : Boning, Forsch. Geb. Pflan- zenkr. u. Immun. Pflanzenreich, J^, 1927, 43-111; Johnson and Jones, Jour. Agr. Res., 54, 1937, 629-638; Loring et al., Proc. Soc. Exp. Biol, and Med., 38, 1938, 239-241; Osborn, Phytopath., 25, 1935, 160-177 ; 28, 1938, 749-754, 923-934 ; Pierce , Jour. Agr. Res., 51, 1935, 1017-1039; Snyder, Phytopath., 24, 1934, 78-80; Stubbs, ihid., 27, 1937, 242-266. 21. Marmor medicaginis H. {loc. cil., 91). From New Latin Medicago, generic name of alfalfa (lucerne). Comnion name : Alfalfa-mosaic virus. Hosts : LEG UMINOSA E— Medicago sativa L., alfalfa (lucerne). SOLAN A- CEAE — Solanum tuberosum L., potato. Experimentally, also transmissible to many species of dicotyledonous plants (summarized by Price, Am. Jour. Bot., 27, 1940, 530-541) including CL^CL' 7?^/- TACEAE — Cucumis sativus L., cucum- ber. COM POS I T AE— Zinnia elegans Jacq., zinnia. LEGUMINOSAE— Phaseolus vulgaris L., bean; Trifolium incarnaium L., crimson clover. SOL- ANACEAE — Capsicum frutescens L., pepper; Lycopersicon esculentum ]\Iill., tomato; Nicotiana tabacum L., tobacco. Geographical distribution : United States. Induced disease : In alfalfa, systemic chlorotic mottling, tending to be masked at times. In bean, (most varieties) small necrotic primary lesions, reddish brown at periphery. No secondary le- sions. Some bean varieties show no lesions after inoculation; one of these, Refugee Rogue, possesses two dominant genes either of which will confer this type jf resistance. In tobacco, white necrotic flecks, small rings and arcs on inoculated leaves ; later, systemic mottling, followed by production of necrotic oak-leaf pat- terns ; virus content may be low in plants long diseased, especially in summer. Transmission : Bj^ inoculation of ex- pressed juice. By aphids, Macrosiphum pisi Kalt. (for typical strain) and M. solanifolii Ashm. (for potato-calico strain) (APHIDIDAE). Not through seeds from diseased plants. Immunological relationships : Resis- tance to superinfection with the type of this virus is conferred by earlier infection with potato-calico virus (now considered a related strain but earlier regarded as distinct), but not by earlier infection with potato-mottle virus, cucumber- mosaic virus, or the Canada-streak strain of potato aucuba-mosaic virus. Thermal inactivation : At 65 to 70° C in 10 minutes. Other properties : Sedimentation con- stant, 73.9 ± 5.2 X 10-13 cm. per sec. in a unit centrifugal field. Specific volume 0.673. Particles spherical or nearly so. Diameter 16.5 millimicrons; weight 2.1 X 10^ times hydrogen unit. Isoelectric point about pH 4.6. Inacti- vated and, more slowly, hydrolyzed by trypsin. Literature : Black and Price, Phyto- path., 30, 1940, 444-447 ; Lauffer and Ross, Jour. Am. Chem. Soc, 62, 1940, 3296-3300, Pierce, Phj^topath., 24, 1934, 87-115; Price, Am. Jour. Bot., 27, 1940, 530-541; Ross, Phytopath., 31, 1941, 394-410, 410- 420; Wade and Zaumeyer, Jour. Am. Soc. Agron., 32, 1940, 127-134; Zaumeyer, Jour. Agr. Res., 56, 1938, 747-772. Strains : At least one strain of alfalfa- mosaic virus was formerly considered as an independent virus, causing a disease known as calico in potato. It has now been given varietal rank and distin- guished from the type, var. typicum Black and Price (Phj^topath., 30, 1940, 446) by the following name : 21a. Marmor medicaginis var. solani Black and Price (Phytopath., 30, 1940, 1182 MANUAL OF DETERMINATIVE BACTERIOLOGY 446). From New Latin Solanum, gen- eric name of potato. Common name : Potato-calico strain of alfalfa-mosaic virus. Differing from the type by inducing a more severe disease in potato, in which it is commonly found in nature. (Price and Black, Phytopath., 30, 1940, 444-447 ; Dykstra, ibid., 29, 1939, 917-933; Porter, Potato Assoc. Amer. Proc, 18, 1931, 65-69; Hilgardia, 6, 1931, 277-294; 9, 1935, 383-394.) 22. Marmor tulipae H. {loc. cit., 52). From New Latin Tulipa, generic name of tulip. Common name : Tulip color-adding virus. Hosts : LILIACEAE — Tulipa gesneri- ana L., garden tulip; T. eichleri Regel ; T. greigi Regel. Insusceptible species: AMARYLLI- DACEAE — Narcissus sp., narcissus. IRIDACEAE — Iris germanica L., iris. LILIACEAE — Allium cepa L., onion. SOLAN ACEAE — Nicotiana tabacum L., tobacco. Geographical distribution : Wherever hybrid tulips are grown. Induced disease : In tulip, no obvious effect on leaves but dark striping of flower by pigment intensification. Little inter- ference with growth of plant. No intra- cellular bodies. Transmission : B}' hypodermic injec- tions of expressed juice in dilutions to 10~*. By aphids, Myzus persicae (Sulz . ) , Macrosiphum solanijolii Ashm. (= M. gei Koch, Illinoia solanijolii Ashin.), Aphis (= Anuraphis) tulipae B. de Fonsc. (on bulbs), and perhaps Macrosi- phum pelargonii Kalt. (APHIDIDAE). Not through seeds from diseased plants. Thermal inactivation : At 65 to 70° C in 10 minutes. Literature : Hughes, Ann. Appl. Biol., 18, 1931, 16-29; 21, 1934, 112-119; Mc- Whorter, Phytopath., 22, 1932, 998 (Abst.); 25, 1935, 898 (Abst.); Ann. Appl. Biol., 25, 1938, 254-270. 23. Marmor mite H. (loc. cit., 53). From Latin rnitis, mild. Common name : Lily latent-mosaic virus. Hosts : LILIACEAE — Lilium amabile; L. auratum Lindl.; L. canadense L.; L. candidum L.; L. cernuum; L. chalcedoni- ciim L. ; L. croceum Chaix. ; L. davmottiae; L. elegans Thunb.; L. formosanum Stapf.; L. giganteum; L. henryi Baker; L. leucanthum; L. longiflorum Thunb.; L. myriophyllum; L. pumilum; L. regale Wils.; L. sargentiae Wils.; L. speciosum Thunb.; L. superbum L.; L. testaceum Lindl.; L. tigrinum Ker; L. umbellatum Hort. ; L. wallacei; Tulipa gesneriana L., garden tulip; T. clusiana Vent.; T. lini- folia Regel. Insusceptible species : LILIACEAE — - Allium cepa L., onion; Lilium hansoni Leichtl. IRIDACEAE — Iris germanica L., iris. SOLAN ACEAE — Nicotiana ta- bacum L., tobacco. Geographical distribution : Wherever lilies and tulips are cultivated. Induced disease : In Easter lily, masked symptoms or systemic chlorotic mottling, in either case without necrotic flecking. In tulip, systemic chlorotic mottling in foliage and flower "breaking" (color re- moval, except in a few varieties in which color intensification occurs instead) . In- tracellular bodies characterize ijivaded tissues. Transmission : By inoculation of ex- pressed juice (rubbing surface of leaves), in both lily and tulip. By plugging and grafting of dormant bulbs of tulip. By aphids, Myzus persicae (Sulz.), Macro- siphum solanifolii Ashm. (= M. gei Koch), and Aphis (= Anuraphis) tulipae B. de Fonsc. (APHIDIDAE). Not through seeds from mosaic Lilium longi- florum . Thermal inactivation: At 65 to 70° C in 10 minutes. Literature: Atanasoff, Bull. Soc. Bot. Bulgarie, 2, 1928, 51-60; Brierley, Phyto- path., 29, 1939, 3 (Abst.); 30, 1940, 250- FAMILY MARMORACEAE 1183 257; 31, 1941, 838-843; Brierley and Doo- little, ibid., 30, 1940, 171-174; Cayley, Ann. Appl. Biol., 15, 1928, 529-539; 19, 1932, 153-172; Guterman, Hort. Soc. N. Y., Yearbk., 1930, 51-102; Hall, Gard. Chron., 93, 1933, 330-331; Hughes, Ann. Appl. Biol,. 21, 1934, 112-119; McKay and Warner, Nat. Hort. Mag., 12, 1933, 179-216; McWhorter, Phytopath., 25, 1935, 898 (Abst.); Science, 86, 1937, 179; Ann. Appl. Biol., 25, 1938, 254-270; Science, 88, 1938, 411; Ogilvie and Guterman, Phytopath., 19, 1929, 311-315. 24. Manner iridis H. (loc. cit., 55). From New Latin Iris, generic name of iris. Common name : Iris-mosaic virus. Hosts: IRIDACEAE—Iris filifolia Boiss., I. tingitana Boiss. and Reut., and I.xiphiuyn L., bulbous irises ; Iris ricardi Hort.; /. ungnicularis Poir.; bearded iris, variety William Mohr. Insusceptible species: SOLAN A- CEAE — Lycopersicon esculentum Mill., tomato; Nicotiana tabacum L., tobacco; Petunia hybrida Vilm., petunia. LILIA- CEAE — Tiilipa gesneriana L., tulip. Geographical distribution : United States (Washington, Oregon, California), Holland, Bulgaria, France, England. Induced disease : In bulbous irises, dwarfing of plant, chlorotic mottling of foliage, breaking of flowers. Rate of increase in planting stock decreased. Flower breaks usually darker than normal color of flower. Vacuolate intracellular bodies in some affected tissues. Transmission : By injection of freshly extracted juice of diseased plants into internodal tissue. By aphids, Macro- siphum (= Illinoia) solanifolii Ashm. and Myzus persicae (Sulz.) (APHIDI- DAE). Literature : Brierley and McWhorter, Jour. Agr. Res., 53, 1936, 621-635. 25. Marmor sacchari H. {loc. cit., 60). From New Latin Saccharum, generic name of sugar cane, from Latin sac- charum, sugar. Common name : Sugar-cane mosaic virus. Hosts : GRAM I NEAE— Saccharum of- ficinarum L., sugar cane ; Holcus sorghum L., sorghum; H. sudanensis Bailey, Sudan grass ; Brachiaria platyphylla Nash ; Chaetochloa magna Scribn. ; C. ver- ticillata Scribn. ; Paspalum boscianum Fluegge ; Syntherisma sanguinale Dulae. E.xperimentally, also Zea mays L., corn (maize); Chaetochloa lutescens Stuntz ; Echinochloa crusgalli Beauv. ; Miscanlhus sinensis Anderss., eulalia; Panicum di- chotomiflorum Michx.; Pennisetum glau- cum R. Br., pearl millet; Saccharum narenga Wall. Insusceptible species : All tested spe- cies other than Gramineae. Geographical distribution : Originally in Far East; now in nearly all countries where sugar cane is grown; believed still to be absent from Mauritius. Induced disease: In sugar cane, sys- temic mottling chlorosis, light areas of pattern elongated, but crossing veins. Occasionally, stem cankers. Regularly, discoloration and necrosis in mature inner stalk tissues. Vacuolate intracellular bodies occur in diseased tissues. Canes sometimes recover, spontaneously losing the virus and becoming susceptible to reinfection. Transmission : By inoculation of ex- pressed juice (puncture through inocu- lum into young leaf). By aphids. Aphis maidis Fitch, Carolinaia cyperi Ainslie, Hysteroncura setariae (Thomas), and Toxoptcra graminum Rond.; not by Sipha flava Forbes (APHIDIDAE). Not by Draeculacephala mollipcs (Say) (CICADELLIDAE). Serological relationships: Specific neu- tralizing and precipitating antibodies have been demonstrated. Thermal inactivation : At 53 to 54° C in 10 minutes in leaf tissues. 1184 MANUAL OF DETERMINATIVE BACTERIOLOGY Other properties : Active after storage 27 days at -6° C. Literature: Brandes, Jour. Agr. Res., 19, 1920, 131-138, 517-522; U, 1923, 247- 262; Desai, Current Science, 3, 1935, 18; Forbes and Mills, Phytopath., 33, 1943, 713-718; Ingram and Summers, Jour. Agr. Res., 5£, 1936, 879-888; Kunkel, Bull. Exp. Sta. Hawaiian Sugar Planters' Assoc, Bot. Ser., 3, 1924, 115-167; Matz, Jour. Agr. Res., 46, 1933, 821-839; Raf ay, Indian Jour. Agr. Science 5, 1935, 663- 670; Sein, Jour. Dept. Agr. Porto Rico, U, 1930, 49-68; Stoneberg, U. S. Dept. Agr., Tech. Bull. 10, 1927; Tate and Vandenberg, Jour. Agr. Res., 59, 1939, 73-79. 26. Marmor cepae H. {loc. cit., 66). From Latin cepa, onion. Common name: Onion yellow-dwarf virus. Host: LILIACEAE — Allium cepa L., onion (the variety viviparum Metz. is symptomless when infected and may serve as an unrecognized reservoir of virus). Geographical distribution : United States, Germany, Czecho-Slovakia, Rus- sia, New Zealand. Induced disease : In onion (most varie- ties), yellow streaks at base of developing leaf, followed by yellowing, crinkling, and flattening of newly formed leaves; leaves prostrate, flower stalks bent, twisted, stunted; plants reduced in size, bulbs small, yield of seeds reduced. A few varieties of onion are relative!}' tolerant, and the tree-onion, var. vivi- parum is symptomless after infection. Transmission: By inoculation of ex- pressed juice. By 48 of 51 tested species of aphid, principally Aphis rum ids Linn., A. maidis Fitch, and lihopalosiphiim prunifoliae Fitch (APHIDIDAE). Not through seeds from diseased plants. Not by contaminated soil. Thermal inactivation : At 75 to 80° C in 10 minutes. Other properties : Virus withstands di- lution to 10~^ storage at 29° C for about 100 hours and storage at — 14° C for more than time tested (6 hours), but is inacti- vated by drying in leaf tissues. Literature : Andreyeff, Rev. Appl. Mycol., 17, 1938, 575-576; Blattny, Och- rana Rostlin, 10, 1930, 130-138; Bremer, Phytopath. Ztschr., 10, 1937, 79-105; Brierley and Smith, Phytopath., 34, 1944, 506-507; Chamberlain and Baylis, New Zealand Jour. Science and Technology, 21, 1939, 229A-236A; Drake et al., Iowa State Coll. Jour. Science, 6, 1932, 347- 355; Jour. Econ. Ent., 26, 1933, 841-846; Henderson, Phytopath., 20, 1930, 115 (Abst.); Iowa State Coll., Research Bull. iSS, 1935, 211-255; Melhus et al., Phytopath., 19, 1929, 73-77; Porter, U. S. Dept. Agr., Plant Dis. Rept., 12, 1928, 93; Tate, Iowa State Coll. Jour. Science, 14, 1940, 267-294. 27. Manner scillearum Smith and Brierley (Phytopath., 34, 1944, 503.) From New Latin Scilleae, name of tribe in which hosts are classed. Common name : Ornithogalum-mosaic virus. Hosts: LILIACEAE (of the tribe Scilleae) — Ornithogalum thyrsoides Jacq. ; probably also Galionia candicans Decne . ; Hyacinthus orienialis L., hyacinth; La- chenalia sp. Insusceptible species : LILIACEAE (of the tribe Scilleae) — Muscari botry- oidcs Mill. ; Scilla peruviana L. ; Camassia leichtlinii (Baker) S. Wats.; Hyacinthus azureus (Fenzl.) Baker. AMARYLLI- DACEAE — Pancratium maritimum; Ze- phyranthus sp. IRIDACEAE—Tri- toniacrocata (L.) Ker. LILIACEAE — Agapanthus africanus; Allium cepa, onion; A. cernuum Roth.; A. fistulosum Ij.; A. porrum L. ; Gloriosa rothschildiana O'Brien; Lilium formosanum Stapf.; and L. longiflorum. SOLAN ACE AE—Ni- cotiana tahacum L. Geographical distribution : United States (Oregon; probably also Alabama and presumed to be widespread in plants of the squill tribe, Scilleae, of the family LILIACEAE). FAAULY MARMORACEAE 1185 Induced disease -.In Ornithogalumthyr- Transmission: By inoculation of ex- soides, young leaves finely mottled with pressed juice in the presence of fine light and dark green, and becoming more carborundum powder, with difficulty, conspicuously mottled with gray or yel- By aphids. Aphis gossypii Glov., Macro- low as the leaves mature; flower stalks siphumliluMone\\,M.solanifolnAshm.., sometimes boldlj' marked with light and and Myzus persicae (Sulz.); less effici- dark green blotches. In perianth seg- ently by Myzus circumflexus (Buckt.) ments, thin longitudinal streaks. (APHIDIDAE). Key to the species of the Miscellaneous Mosaic-Virus Group. Many of the following viruses, although described in some detail in the literature, stand in need of reinvestigation to determine additional properties and possible rela- tionships to preceding groups. I. Affecting species of MALVACEAE. 28. Marmor abutilon. II. Affecting species of CELASTRACEAE. 29. Marmor euonymi. III. Affecting species of OLE ACE AE. 30. Marmor ligustri. IV. Affecting species of LEGUMINOSAE (and no. 39, other families also). 31. Marmor laburni. 32. Marmor arachidis. 33. Marmor trifolii. 34. Marmor pachyrhizi. 35. Marmor vignae. 36. Marmor repens. 37. Marmor Jastidiens. 38. Marmor iners. 39. Marmor efficiens. V. Affecting species of GRAMINEAE. 40. Marmor tritici. 41. JM armor graminis. VI. Affecting species of MUSACEAE. 42. Marmor abaca. VII. Affecting species of PASSIFLORACEAE. 43. Marmor passiflorae. VIII. Affecting species of ROSACE AE. 44. Marmor flaccumfaciens. 45. Marmor rosae. 46. Marmor veneniferum. 47. Marmor mali. 48. Marmor jragariae. 49. Marmor marginans . 50. Mar7nor rubi. 51. Marmor persicae. 52. Marmor astri. 53. Marmor rubiginosum. 54. Marmor cerasi. 55. Marmor lineopictum. 56. Marmor pallidolimbatus . 57. Marmor nerviclarens. 1186 MANUAL OF DETERMINATIVE BACTERIOLOGY IX. AEecting species oiV IT AC EAE. 58. Marmor viticola. X. Affecting species of SANTALACEAE. 59. Marmor santali. XI. Affecting species of CONVOLVULACEAE and, experimentally, of other families. 60. Marmor secretum. XII. Affecting species of GERANIACEAE. 61. Marmor pelargonii. XIII. Affecting species of SOLA N ACE AE and in most cases also of other families. 62. Marmor angliae. 63. Marmor aevi. 64. Marmor raphani. XIV. Affecting species of PRIMULACEAE. 65. Marmor primulae. XV. Affecting species of MORACEAE. 66. Marmor caricae. XVI. Affecting species of RUT ACE AE. 67. Marmor italicum. 28. Marmor abutilon H. {loc. cit., 50). From New Latin Abutilon, generic name of a host . Common name : Abutilon-mosaic virus. Hosts: M ALV ACE AE— Abutilon stri- atum Dicks, var. thompsonii Veitch. Experimentally, also Abutilon arboreum Sweet; A. avicennae Gaertn.; A. esculen- tum St. Hil.; A. indicum Sweet; A. in- signe Planch. ; A . megapotamicuTn St. Hil. and Naud. ; A. regnellii Miq. ; A. sellowi- anum Regel; A. venosum Lem.; A. viti- folium Presl.; Althaea ficifolia Cav.; A. officinalis h.; A. rosea Cav.; Anoda has- tata Cav.; Kitaibelia vitifolia Willd.; Malva borealis; M . crispa; M. mauritiana Mill. ; M. sylvestris L. ; M. verticillata L. ; Malvastrum capense Garcke; Modiola de- cumbens G. Don.; Sida mollis Herb.; S. napaea Cav.; Sidalcea Candida A. Gray. Insusceptible species: MALVACEAE — Althaea taurinensis; Sidalcea purpurea; Sphaeralcea umbellata G. Don. Geographical distribution : Germany, France, England, United States; orig- inally obtained from a single variegated seedling found among green plants of Abutilon striatum imported from the West Indies in 1868 by Veitch and Sons ; subsequently the infected plant was propagated vegetatively as an ornamental variety. Induced disease : In Abutilon, systemic chlorotic mottling. Recovery occurs if there is persistent removal of affected leaves, suggesting that the virus does not increase in stems. After recovery, plants are susceptible to reinfection. Transmission : By grafting, except patch-bark-grafting, which is ineffective. Occasionally through seeds from diseased plants. Not by inoculation of expressed juice. No insect vector is known. Varieties : Distinctive strains have been noted, but not separately named; one isolate originally occurring in Abuti- lon darwini var. tesselatum, seems to be- long here ; it differs from the type princi- pally in severity of induced disease and in ability to infect Lavatera arborea. Literature : Baur, Ber. d. Deutsch. Bot. Gesellsch., 22, 1904, 453-460; 24, 1906, 416-428; 25, 1907, 410-413; K. Preuss. Akad. AViss., Sitzungsber., 1906, 11-19; Davis, Ann. Missouri Bot. Gard., 16, 1929, 145-226; Hertzsch, Ztschr. f. Bot., 20, 1927, 65-85; Keur, Phytopath., 23, 1933, 20 (Abst.) ; 2^, 1934, 12-13 (Abst.) ; Bull. Torrey Bot. Club, 61, 1934, 53-70; Lindemuth, Gartenflora, 51, 1902, 323- 326. FAMILY MARMORACEAE 1187 29. Marmor euonymi H. (loc. cit., 51). From New Latin Eiwnymus , generic name of host. Common name : Euonymus-mosaic virus. Hosts: CELAST RACE AE— Eiwny- mus japonica L. f. (sometimes written Evonymus japonicus) . Probably also E. radicans Sieb. Geographical distribution: Germany. Induced disease: In Euonymus japon- ica, persistent yellowing along veins. Transmission :.By grafting. Literature: Baur, Ber. d. Deutsch. Bot. Gesellsch., 26a, 1908, 711-713; Rischkow, Biol. Zentralbl., ^7 , 1927, 752-764. 30. Marmor ligustri H. {loc. cit., 52). From New Latin Ligustrum, generic name of host, from Latin ligiistriim, ancient name of privet plant. Common name : Ligustrum-mosaic virus. Host : OLE ACE AE — Ligustrum vul- gar e L., common privet. Geographical distribution: Germany. Induced disease : Systemic chlorotic spotting. Transmission : By grafting. Not through seeds from diseased plants. Literature: Baur, Ber. d. Deutsch. Bot. Gesellsch., 25, 1907, 410-413. 31. Marmor laburni H. {loc. cit., 51). From generic name of a host plant. Laburnum vulgare. Common name : Laburnum-mosaic virus. Hosts : LEG UM I NOSAE— Laburnum vulgare Griseb. {= L. anagyroides Med- ic), bean tree. Experimentally, also Cytisus hirsutus L. Insusceptible species: LEGUMINO- SAE — Laburnum alpinum Griseb.; Cy- tisus purpureus. Geographical distribution: Germany. Induced disease : Systemic chlorotic variegation. Transmission : By bark grafts or by budding. Not through seeds from dis- eased plants of Laburnum vulgare. Literature: Baur, Ber. d. Deutsch. Bot. Gesellsch., 25, 1907, 410-413. 32. Marmor arachidis H. {loc. cit., Q7). From New Latin Arachis, generic name of peanut . Common name: Peanut-rosette virus. Host : LEGUMINOSAE— Arachis hy- pogaea L., peanut. Geographical distribution : Union of South Africa, Madagascar, Tanganyika Territory, L'ganda, Senegal, Gambia, Sierra Leone, Java. Induced disease: In peanut, yellowing of young leaves, at first with green veins ; reduction in leaf size, petiole length, and internode length, producing rosette ; curling and distortion of later-formed, wholly chlorotic or chlorotically mottled leaflets. Seed formation inhibited. No abnormal proliferation of tissues. Transmission : By grafting. By both winged and wingless individuals of the aphid. Aphis laburni Kalt. {= A. legum- inosae Theob.) {APHIDIDAE). Not by 13 tested species of leafhoppers. Not by inoculation of expressed juice. Not through seed from diseased plants. Not through soil. Literature : Hayes, Trop. Agr., 9, 1932, 211-217; McClintock, Science, 45, 1917, 47-48; Soyer, Publ. Inst. Nat. Etud. Agron. Congo Beige, Ser. Sci., 21, 1939, 23 pages (Rev. Appl. Mycol., 19, 1940, 386, Abst.) ; Storey and Bottomley, Ann. Appl. Biol., 15, 1928, 26-45; Zimmerman, Der Pflanzer, 3, 1907, 129-133; 9, 1913, 59-63. 33. Marmor trifolii H. {loc. cit., 93). From New Latin Trijoliimi, generic name of red clover, from Latin trifolium, clover. Common name : Red-clover vein-mo- saic virus. Hosts : LEGUM I NOSAE— Trifolium pratense L., red clover; Lathyrus odoratus L., sweet pea; Viciafaba L., broad bean. Experimentally, also Trifolium hybridum L., alsike clover; T. incarnatum L., crim- 1188 MANUAL OF DETERMINATIVE BACTERIOLOGY son clover; T. repens L., white clover; Melilotvs alba Desr., white sweet clover; Pisum sativum L., pea. Insusceptible species: LEGUMINO- SAE — Phaseolus vulgaris L., bean; P. aureus Roxb., mung bean; Medicago sali- va L., alfalfa. SOLAN ACE AE—Lyco- persicon esculentum Mill., tomato; Nico- tiana lahacum L., tobacco; N. glutinosa L.; N. langsdorffii Weinm.; A^. rustica L.; N. sylvestris Spegaz. and Comes; Solanum tuberosum L., potato. Geographical distribution : United States. Induced disease : In red clover, yellow color along veins, but no mottling. Some- times small yellow spots in interveinal areas. Little or no stunting. In Vicia faba, experimentally, necrotic splotches or rings sometimes at site of inoculation. Clearing of veins followed by appear- ance of whitish bands along the veins. Stalks discolored, purplish. Diseased plants are stunted and often die back to a point near the base of the stalk, inducing new growth from buds on the stem. Transmission : By inoculation of ex- pressed juice, using carborundum. By aphid, Macrosiphum pisi Kalt. (APHP DIDAE), without incubation period and without long retention. Not by aphids, Macrosiphum solanijolii Ashm. (= M. gei Koch) or Aphis rumicis Linn. (APHIDIDAE). Thermal inactivation : At 60° C in 10 minutes. Literature: Osborn, Phytopath., 27, 1937, 1051-1058; Zaumeyer, Jour. Agr. Res., S6, 1938; 747-772; Zaumeyer and Wade, Phytopath., 27, 1937, 1009-1013. 34. Marmor pachyrhizi spec. nov. From New Latin Pachyrhizus, generic name of sincamas. Common name : Sincamas-mosaic virus. Host : . LEG UM I NOSAE— Pachyrhi- zus erosus (L.) Urb., sincamas (yam bean). Insusceptible species: LEGUMINO- SAE — Phaseolus vulgaris L., bean. Geographical distribution : Philippine Islands. Induced disease : In sincamas, chlorotic mottling of foliage; in plants infected when young, dwarfing. Transmission: By inoculation of ex- pressed juice, in the presence of sand as abrasive. Through about 25 percent of the seeds from infected plants. Not through soil, interlacing of roots, or casual contacts of leaves and stems. No insect vector is known. Literature: Fajardo and Maranon, Philippine Jour. Science, 48, 1932, 129- 142. 35. Marmor vignae spec. nov. From New Latin Vigna, generic name of cow- pea, from family name of an Italian botanist, Domenico Vigna. Common name : Cowpea-mosaic virus. Hosts : LEGUM I NOSAE— Vigna sin- ensis (L.) Endl., cowpea. Experimen- tally, also Phaseolus lunatus L., lima bean. Geographical distribution: United States (Arkansas, Oklahoma, Louisiana, Indiana, Georgia, Iowa, Mississippi, Kan- sas, New Jersey). Induced disease : In cowpea, clearing of veins followed by chlorotic mottling, slight convex cupping of leaflets, short- ened internodes, abortion of flowers, twisting of petioles, delayed maturity. Malformation of leaves, stunting of plants, and reduction of yield more pro- nounced in some varieties of cowpea than in others. Transmission : By inoculation of ex- pressed juice, especially in the presence of fine carborundum powder. By aphids, Macrosiphum solanifolii Ashm., M. pisi Kalt., Aphis gossypii Glov. {APHIDI- DAE) ; not by various beetles nor by the bean leaf hopper, Empoasca fabae LeB. (CICADELLIDAE). Through 5 per cent of seeds from infected cowpea plants. Thermal inactivation: At 72 to 75° C in 10 minutes. Other properties : Infectious in dilu- tions as high as 1 :1000 and after 2 days FAMILY aiAKMORACEAE 1189 storage in expressed juice at room tem- perature, 20 to 25° C. Literature: Elliott, Phytopath., 11, 1921, 146-148; Gardner, Indiana Acad. Science Proc, 36, 1927, 231-247; 37, 1928, 417; McLean, Phytopath., 31, 1941, 420- 430; Smith, Science, 60, 1924, 268. 36. Marmor repens Johnson. (Phyto- path., 32, 1942, 114.) From Latin repens, unlooked for, in reference to unexpected discovery of this virus as a constituent of a complex formerly regarded as a single virus, so-called "white-clover mosaic virus". Common name : Pea-wilt virus. Hosts : LEG UMINOSAE—Trifolium repens L., white clover. Experimen- tally, also Lathyrus odoratus L., Lens escu- lenia Moench.; Lupinus albns L. ; Medi- cago lupulina L. ; Melilotus alba Desr.; Phaseolvs aureus Roxb., mung bean; P. vulgaris L., bean; Pisum sativum L., pea; Trijolium hybridum L.; T. incarnatum L.; T. praiense L.; Vicia faba L.; V. saliva L.; Vigna sinensis (L.) Endl., cowpea. Insusceptible species : CARYOPH YL- LACEAEStellaria media (L.) Cyrill. C HENOPODI ACE AE— Beta vulgaris L.; Spinacia oleracea L. COMPOSI- TAE — Callistephus chinensis Nees; Lac- tuca saliva L.; Taraxacum officinale Weber; Zinnia elegans Jacq. CRUCI- FERAE — Barbarea vulgaris R. Br.; Brassica oleracea L.; Raphanus sativus L. CUCURBITACEAE—Cucumis sa^ tivus L. GRAMINEAE—Zea mays L. LEGUM I NOSAE— Glycine max Merr.; Lupinus hirsutus L.; Medicago saliva L. LILIACEAE — Lilium formosanum Stapf. PLANTAGINACEAE—Plan- tago lanceolata L.; P. major L. POLY- GON ACEAE — Rumex acetosella L. SCROPHULARI ACEAE — Antirrhi- num majus L. SOLAN ACEAE — Da- tura stramonium L.; Lycopersicon escu- lentum Mill.; Nicotiana glutinosa L.; N. rustica L.; N. sylvestris Spegaz. and Comes; N. labacum L.; Solanum nig- rum L. Geographical distribution : United States (Washington). Induced disease: In white clover, sys- temic chlorotic mottling. In pea, experi- mentally, originally infected leaves wilt and die, remaining attached to the stem by their shriveled petioles ; a few adjacent lower leaves may also wilt and die; in most varieties the top foliage remains green, but in two varieties, Alaska and Canada White, it mottles faintly; stems show faint grayish discoloration; plants are retarded in growth and dwarfed. If pea-mottle virus, Marmor efficiens John- son, is also present, a severe streak dis- ease occurs. Intracellular inclusions ab- sent. In mung bean, experimentally, necrotic zonate local lesions. In cowpea, experimentally, brown necrotic local lesions in inoculated primary leaves, diffuse areas of bleaching in uninoculated trifoliate leaves. In bean, experimen- tally, mild chlorotic mottling except in three varieties that appear insusceptible (varieties Ideal Market, Kentucky Won- der, and Navy Robust). Transmission : By inoculation of ex- pressed juice. Not by dodder, Cuscuta campestris Yunck. {CONVOLVULA- CEAE). Not by pea aphid, Macro- siphum pisi Kalt. (APHIDIDAE). No insect vector is known. Thermal inactivation : At 58 to 60° C in 10 minutes. Filterability : Passes Berkefeld W filter candle. Other properties : Infectious in dilution of 1 : 100,000. Not inactivated by storage in juice of infected plants at about 25° C for one month or by similar storage in dried tissues of infected pea plants. Literature: Johnson, Phytopath., 32, 1942, 103-116; Pierce, Jour. Agr. Res., 51, 1935, 1017-1039. 37. Marmor fastidiens spec. nov. From Latin fastidiens, disdaining, in reference to slight irregularities in the reported host ranges of constituent strains and failure of this virus to infect certain varieties of the pea although it 1190 MANUAL OF DETERMINATIVE BACTERIOLOGY may utilize many other varieties of this species as host. Common name: Alsike-clover mosaic virus. Hosts : LEGUM I NOSAE—Trifoliiwi hybridum L., alsike clover; Pisum sati- vum L., pea (except the varieties Horal, Perfection, and Surprise). Experimen- tally, also Crotalaria striata DC; C. retusa L.; and C. spectabilis Roth (the two last-named species are reported to be insusceptible to the type strain of the virus, but susceptible to one or more of the other tested strains) ; Lupintis albus L. ; L. angvstifolius L.; Medicago saliva L.; Melilotus alba Desr.; Phaseolus vul- garis L., bean; Trifolium incarnatum L.; T. pratense L.; Vicia faba L. Insusceptible species: SOLAN A- CEAE — Datura stramonium L. ; Nico- tiana glauca Graham; N. glutinosa L.; A^. tabacujn L. ; Petiinia hybrida Vilm. LEG UM I NOSAE— Phaseolus aureus Roxb., mung bean; P. lunatus L., sieva bean; Soja max (L.) Piper, soybean; Trijoliuvi repens L., white clover; Vicia saliva L., spring vetch. Induced disease : In pea and bean, ex- perimentally, systemic chlorotic mot- tling; some isolates kill inoculated leaves and even cause death of infected plants. Transmission : By inoculation with ex- pressed juice, at dilutions to 1:6000 or 1 : 8000. No insect vector is known. Thermal inactivation : At 60 to 65° C in 10 minutes; one strain at lower tempera- ture, 54 to 58° C. Strains : Several strains have been dis- tinguished by the severity of their effects on host plants. These may be charac- terized as follows: var. fastidiens, var. nov., type variety, the first of the strains to be described (originally known as alsike clover mosaic virus 1 ), induces mild disease in pea, does not infect red clover ; var. viite, var. nov., described as pea mosaic virus 4, induces mild symptoms on pea, infects red clover; var. reprimens , var. nov., described as pea mosaic virus 5, stunts peas severely; var. denudans, var. nov., described as alsike clover mosaic virus 2, defoliates pea plants. Varietal names from New hat'in fastidiens , epithet of the species, and from Latin milis, mild ; reprimere, to restrain; and denudare, to denude ; all three in reference to induced symptoms. Literature: Wade and Zaumeyer, Phy- topath., 28, 1938, 505-511; Zaumeyer, Jour. Agr. Res., 60, 1940, 433-452. 38. Marmor iners spec. nov. From Latin iners, sluggish or inert, in reference to failure of the virus to spread systemi- cally in certain of its hosts. ('ommon name : Pea-streak virus. Hosts: LEGUM I NOSAE— Pisum sa- tivum L., pea. Experimentally, also Galega officinalis L., goat's rue; Glycine soja Sieb. and Zucc, soyabean; Lathyrus odoratus L., sweet pea; Lotus hispidus Desf.; Lupinus angustifolius L., blue lupin; L. lutevs L., yellow lupin; L. mutabilis Sweet; Phaseolus vulgaris L., bean; Trifolium arvense L., haresfoot trefoil ; T. cernuum Brot . , nodding clover ; T. fragiferum L., strawberry clover; T. glomeratum L., cluster clover; T. hybri- dum L., alsike clover; T. pratense L., red clover; T. repens L., white clover; Vicia villosa Roth., hairy vetch. CUCUR- BITACEAE — Cucumis melo L., rock melon; C. sativus L., cucumber; Cucur- bita pepo L., marrow. Insusceptible species : CHENOPODI- ACEAE — Spinacia oleracea L., spinach; Beta vulgaris L., beet. COMPOS I TAE — Calendula officinalis L., calendula; Lactuca saliva L., lettuce ; Zinnia elegans Jacq., zinnia. CRUCIFERAE—Brass- ica napus L., swede; B. oleracea L., cab- bage; B. rapa L., turnip; Matthiola in- cana R. Br., stock; Raphayms sativus L., radish; Sisymbrium officinale (L.) Scop., hedge mustard. LEGUM I NOSAE— Arachis hypogaea L., peanut; Lathyrus latif alius L., perennial sweet pea; L. pubescens Hook, and Arn., Argentine sweet pea ; Lotus corniculatus L. ; Lupinus arboreus Sims, tree lupin; Medicago arabica Huds. ; M. sativa L., lucerne (al- falfa) ; Phaseolus multiflorus Willd., run- FAMILY MARMORACEAE 1191 ner bean; Trifolium striahim L., striated clover; T . subterraneuni L., subterranean clover; Vicia faba L., broad bean. PLANT AGIN ACE AE — Plantago lanceolata L., plantain. SCROPHU- LARIACEAE — Antirrhinum majus L. SOLA NACEAE — Cyphomandra betacea Sendt., tree tomato; Datura stramonium L., Jimson weed; Nicotiana glauca R. Grab.; N. rustica L., Turkestan tobacco; N. tabacum L., tobacco; Physalis peru- viana L., Cape gooseberry; Solanum nigrum L. , black nightshade . TROPAE- OLACEAE — Tropaeolum majus L., nas- t urtium. UM BELLI FERAE—Apinm graveolens L., celery. Geographical distribution: Xew Zea- land. Induced disease : In the pea, stunting, wilting of young leaves, purple or purple - brown spotting on young leaves, dark streak on stem. Near tip, stem may die. Stem becomes brittle, tip bent to one side. Pods maj' remain flat and turn dark purple or purple -brown, or if already formed may show purple or purple -brown markings. Older leaves turn yellow, then brown and shrivelled. Infected plants usually die within two or three weeks. In inoculated plants small brown primary lesions, rapidly increasing in size especially along veins, eventually involve the whole leaf; petiole and stem streak follows. Among garden peas, the varie- ties Pride of the Market, Little IVIarvel, Wm. Massey and Autocrat are little af- fected; among field peas, the varieties Unica and White Ivory are equally resis- tant. In cucumber, experimentally, numerous brown, necrotic local lesions, each with light colored center and sur- rounding light -yellow halo. In bean, experimentally, local and systemic necro- sis, stem streak, death of plant. Transmission: By inoculation of ex- pressed juice, best with an abrasive pow- der such as fine sand. Not by Myzus persicae (Sulz.), Macrosiphum solani (APHIDIDAE), nor Thrips tabaci Lind. (THRIPIDAE). Xo insect vec- tor is known. Thermal inactivation : At 78 to 80° C in 10 minutes. Filterability : Passes ^Nlandler filters of preliminarj^, regular, and fine grades. Other properties : Dilution end point 1:10*. Xot inactivated at room tem- perature in 41 days. Literature: Chamberlain, Xew Zea- land Jour. Science and Technology, 20, 1939, 365A-381A. 39. Marmor efficiens Johnson. (Phy- topath., 32, 1942, 114.) From Latin efficiens, effective, in reference to ability of this virus to cause mottling in all tested varieties of pea in contrast with inability of pea-wilt virus, a second constituent of the complex earlier known as "white- clover mosaic virus," to produce such chlorotic symptoms in tested varieties other than Alaska and Canada White. Common name : Pea-mottle virus. Host s : LEG UMINOSAE— Trifolium. repens L., white clover; Pisum sativum L., pea. Experimentally, also CARYO- PHYLLACEAEStellaria media (L.) Cyrill. CHENOPODIACEAE—Spin- acia oleracea L., spinach. CUCURBI- TACEAE^Cucumis sativus L. LE- GUMINOSAE — Lathyrus odoratus L.; Lens esculenta Moench.; Lupinus albus L. ; L. hirsutus L. ; Medicago lupulina L. ; M. saliva L., alfalfa (lucerne); Meliloius alba Desr. ; Phaseolus aureus Roxb. ; P. vulgaris L., bean; Trifolium hybridum L. ; T. incarnalum L.; T. pratense L.; Vicia faba L. ; V. sativa L. SCRO- PH ULARI ACE AE— Antirrhinum ma- jus L. Insusceptible species: CHEXOPODI- ACEAE — Beta vulgaris L., sugar beet. COM POS I T AE — Callistephus chinensis X'ees; Lactuca sativa L.; Taraxacum offi- cinale Weber; Zinnia elegans Jacq. CRUCIFERAE—Barbarea vulgaris R. Br. ; Brassica oleracea L. ; Raphanus sativus L. GRAMINEAE — Zea mays L. LEGUM I NOSAE— Glycine max Merr.; Vigna sinensis (L.) Endl. LILI- ACEAE — Lilium formosanum Stapf. PLANT AGIN ACE AE — Plantago 1192 MANUAL OF DETERMINATIVE BACTERIOLOGY lanceolata L. ; P. major L. POLYGON - ACEAE — Rumex acetosella L. SOLA X- ACEAE — Datura stramonium L. ; Lyco- persicon esculentum Mill. ; Nicotiana glutinosa L. ; N. rustica L. ; N. sylvestris Spegaz. and Comes; N. tabacum L. ; Solanum nigrum L. Geographical distribution : United States (Washington). Induced disease : Experimentally, in pea, developing leaves late in opening; clearing of veins, chlorotic spotting, stunting, chlorotic mottling; stipules mottled; stems, pods, and seeds appear normal. If pea-wilt virus {Marmor re- pens Johnson) is also present, a severe streak disease occurs. Intracellular in- clusions absent. In bean, light yellow spots and clearing of veins. In spinach, severe chlorotic mottling, dwarfing. In alfalfa, streaks of yellowing along veins, chlorotic mottling. Transmission : By inoculation of ex- pressed juice. By dodder, Cuscuta cam- pestrisYunck. {CON VOLVU L ACEAE). Not by pea aphid, Macrosiphum pisi Kalt. (APHIDIDAE). No insect vec- tor is known. Thermal inactivation : At 60 to 62° C in 10 minutes. Filterability : Passes Berkefeld W filter candle. Other properties : Infectious in dilu- tion of 1:10,000 and after storage in ex- pressed juice or dried tissues for one month at about 25° C. Literature: Johnson, Phytopatli., 32, 1942, 103-116; Johnson and Jones, Jour. Agr. Res., 54, 1937, 629-638; Pierce, ibid., 51, 1935, 1017-1039 ; Zaumeyer and Wade, ibid., 51, 1935, 715-749. 40. Marmor tritici H. {lac. cit., 61). From Latin triticum, wheat. Common names : Wheat -mosaic virus, wheat-rosette virus. Hosts: GRAM I NEAE— Triticum aes- tivum L., wheat; Secale cereale L., rye. Experimentally, also all tested species of the tribe Hordeae ; Triticum compactum Host ; T. turgidum L. ; T. durum Desf . ; T. dicoccum Schrank; T. spelta L. ; T. polonicum L. ; T. monococcum L., Hor- deum vulgar e L., barley. Insusceptible species : GRAMINEAE — Bromus inermis Leyss., awnless brome- grass (of the tribe Festuceae). Geographical distribution : United States, Japan. Induced disease: In wheat, systemic chlorotic mottling, with dwarfing in some varieties; vacuolate, rounded intracellu- lar bodies in diseased cells, usually close to nucleus. Some selections of Harvest Queen wheat are resistant. Transmission : Through soil ; remains infectious in soil 6 or more years. By in- oculation of expres.ied juice (needle punc- tures in stem). Not through seeds or stubble of diseased plants. No insect vector is known. Thermal inactivation : Contaminated soil becomes incapable of infecting wheat plants if heated for 10 minutes at 60° C though not if heated for the same length of time at 50° C. Literature : Johnson, Science, 95, 1942, 610; McKinney, Jour. Agr. Res., 23, 1923, 771-800; U. S. Dept. Agr., Bull. 1361, 1925; U. S. Dept. Agr., Circ. 442, 1937; Jour. Agr. Res., 40, 1930, 547-556; Mc- Kinney et al., ibid., 26, 1923, 605-608; Wada and Hukano, Agr. and Hort., 9, 1934, 1778-1790 (Rev. Appl.Mycol., i.^, 1935, 618, Abst.); Jour. Imp. Agr. Exp. Sta., 3, 1937, 93-128 (Rev. Appl. Mycol., 16, 1937, 665, Abst.); Webb, Jour. Agr. Res., 35, 1927, 587-614; 5ff, 1928, 53-75. 41. Marmor graminis McKinney. (Jour. Washington Acad. Sci., 34, 1944, 325.) From Latin gramen, grass. Common name : Bro me -grass mosaic virus. Hosts: GRAMINEAE— Bromus iner- mis Leyss., awnless brome-grass. Ex- perimentally, also Triticum aestivum L., wheat ; Avena sativa L., oat. Geographical distribution : United States (Kansas). Induced disease : In awnless brome- grass, systemic chlorotic mottling of the FAMILY MARMORACEAE 1193 tj-pe cominoiily called yellow mosaic be- cause of the distinctly yellow color of the chlorotic areas in affected leaves. Transmission : By inoculation of ex- pressed juice or of aqueous suspensions of dried diseased tissues; not inactivated by drj'ing in diseased tissues for at least 51 days. Xo insect vector is known. Literature: McKinney et al., Phyto- path., 32, 1942, 331. 42. Marmor abaca H. {loc. cit., 63). P>om common name of host plant . Common name : Abacd bunchy-top virus. Host: MUSACEAE—Musa. textilis Nee, abaca (Manila hemp plant). Insusceptible species : MUSACEAE — Musa sapientum L. vars. cincrea (Blanco) Teodoro, co?npressa (Blanco) Teodoro, lacatan (Blanco) Teodoro, and suaveolens (Blanco) Teodoro; 71/. cavcndisJdi Lamb. Geographical distribution: Philippine Islands. Induced disease : In abacd (Manila hemp plant), chlorotic lines and spots along veins of young leaves, followed by growth of distorted leaves, successively shorter, narrower, stiffer, and more curled along their margins. The green areas of mottled leaves, petioles, and leaf sheaths are darker than normal. Newly formed diseased leaves unfurl earh', but lire short, producing the bunchy top that is referred to in the common name of the disease. Transmission : By the aphid, Penla- lonia nigronervosa Coq. (APHIDI- DAE), vector also of the apparentlj^ dis- tinct banana bunchy-top virus of Aus- tralia. The incubation period of abaca bunchy-top virus in this aphid is between 24 and 48 hours in length. The j)rogeny of viruliferous aphids do not receive the virus directly, but must feed on diseased plants before they can infect healthy abacd. Transmission by inoculation of expressed juice has not been demon- strated. No soil transmission. Literature: Ocfemia, Am. .Jour. Bot., 17, 1930, 1-18; Philippine Agriculturist, 32, 1934, 567-581. 43. Marmor passiflorae H. {loc. cit., 77) . From New Latin Passi flora, generic name of passion fruit. Conmion name: Passion-fruit woodi- ness virus. Hosts : PASSIFLORACEAE—Passi- flora edulis Sims, passion fruit; P. coeru- lea L. Experimentally, also P. alba Link and Otto. Insusceptible species: SOLAN A- CEAE — Datura siramoniuvi L. ; Lycoper- sicon escnlentxim Mill., tomato ; Nicoliana gluiinosa L. ; A'', tabacum L., tobacco. Geographical distribution : Australia (New South Wales, Queensland, Victoria) , Kenja. Induced disease : In passion fruit, growth checked ; leaves puckered, slightly chlorotic or obscurely mottled, curled, twisted, deformed. Clearing of veins has been observed. Color of stems darker green than normal in some places. Fruits short or deformed, discolored, surface sometimes roughened by cracks; so hard as not to be cut through readily. Peri- carp or rind of fruit abnormally thick. Pulp deficient, color deepened. At tem- peratures below 80° F, some abscission of young chlorotic leaves; above 85° F, masking of the disease in most plants. Transmission : By inserting cotton in stem wound after soaking it in expressed juice of diseased plant. By aphids, My- zus persicae (Sulz.), Macrosiphum solani- folii Ashm., and two dark-colored species of the genus Aphis (APTIIDIDAE). Literature : Cobb, Agr. Gaz. New South Wales, 12, 1901, 407-418; Noble, Jour, and Proc. Roj'. Soc. New South Wales, 62, 1928, 79-98; Noble and Noble, ibid., 72, 1939, 293-317; Sinmionds, Queensland Agr. Jour., 45, 1936, 322-330. 44. Marmor flaccumfaciens H. {loc. cit., 73). From hatm flacczis , flabby, and facere, to make. Common names : Rose-wilt virus, rose die back virus. 1194 MANUAL OF DETERMINATIVE BACTERIOLOGY Hosts: ROS ACE AE— Rosa hybrids, roses. Geographical distribution : Australia, especially Victoria; New Zealand; possi- bly Italy. Induced disease : In rose, leaflets crowd ed, brittle, recurved. Defoliation pro- gresses from tip to base of plant. Tips of branches discolor and die back an inch or two. Stem darkens at base. Buds re- main green and begin development, but growth is soon checked by necrosis at tips. Plant may recover temporarily, but not permanently. Transmission : By inoculation of ex- pressed juice (needle-puncture and scratch methods). No insect vector is known. Filterability : Passes Seitz filter (Seitz EK Schichten type, size 6). Literature: Gigante, Boll. Staz. Pat. Veg. Roma, n. s. 16, 1936, 76-94; Grieve, Austral. Jour. Exp. Biol, and Med. Science, 8, 1931, 107-121; Jour. Dept. Agr. Victoria, 1932 and 1933, pages 30-32. 45. Marmor rosae H. {loc. cit., 74). From Latin rosa, rose. Common name: Rose -mosaic virus. Hosts : ROSACE AE— Rosa rugosa Thunb. ; R. chinensis Jacq. var. manetti Dipp.; R. vxidtiflora Thunb.; R. odorata Sweet, tea rose; R. gymnocarpa; Rnbiis parviflorus Nutt. Geographical distribution : United States, England, Bulgaria, Brazil. Induced disease : In Rosa rugosa and R,. chinensis var. manetti, systemic chlorotie mottling. Transmission : By budding and other forms of graftage. Not by inoculation of expressed juice. No insect vector is known. Literature : Baker and Tiiomas, Phyto- path., 32, 1942, 321-326; Brierley, Phyto- path., 25, 1935, 8 (Abst.); Brierley and Smith, Am. Nurseryman, 72, 1940, 5-8; Jour. Agr. Res. , 61 , 1940, 625-660 ; Kramer, Revista de Agricultura, 15, 1940, 301-311 ; O Biologico, 6, 1940, 365-368; McWhorter, U. S. Dept. Agr., Plant Dis. Rep., 15, 1931, 1-3; Milbrath, West. Florist, 13, 1930, 29-30; Nelson, Phytopath., 20, 1930, 130 (Abst.) ; Newton, Rep. Domin. Bot., 1930, Div. Bot., Canad. Dept. Agr., 1931, 23; Thomas and Massey, Hilgardia, 12, 1939, 645-663; Vibert, Jour. Soc. Imp. et Cent. Hort., 9, 1863, 144-145; White, Phytopath., 22, 1932, 53-69; 24, 1934, 1124-1125. 46. Marmor veneniferum H. (loc. cit., 75). From Latin venenifcr, poisonous, in reference to occasional killing of tissues near inserted bud in graft transmission. Common name: Rose-streak virus. Hosts: ROSACEAE—Rosa multi flora Thunb.; R. odorata Sweet; 72osa hybrids. Geographical distribution: Eastern United States. Induced disease : In various rose species and hybrids, brownish or reddish ring and veinbanding patterns on leaves, and ring patterns on stems. Sometimes necrotic areas near inserted bud, causing girdling of stem and wilting of foliage. Transmission : By grafting. Not by inoculation of expressed juice. No insect vector is known. Literature: Brierley, Phytopath., 25, 1935, 7-8 (Abst.); Brierley and Smith, Jour. Agr. Res., 61, 1940, 625-660. 47. Marmor mail H. (loc. cit., 75). From Latin mains, apple tree. Common name : Apple-mosaic virus. Hosts: ROSACEAE—Pyrus jnalus L., apple. Experimentally, also Cotoneaster harroviana; Eriohotrya japonica Lindl., loquat ; Photinia arbutifolia Lindl., toyon; Rosa sp., rose; Sorbus pallescens. Insusceptible species: ROSACEAE — Amelanchier alnifolia Nutt.; Crataegus douglasii Lindl.; Pyrus communis L., pear. Geographical distribution : United States, Australia, Bulgaria, British Isles. Induced disease : In apple, clearing of veins and systemic chlorotie spotting. The chlorotie areas sometimes become necrotic during months of intense sun- light. Transmission : By grafting. No insect FAMILY MARMORACEAE 1195 vector is known. Transmission by in- oculation of expressed juice has not been demonstrated. Thermal inactivation : Not demon- strated. Virus in stem tissues with- stands at least 50° C for as much as 60 minutes without being inactivated. Literature: Blodgett, Phytopath., 28, 1938, 937-938; Bradford and .loley, Jour., Agr. Res., 46', 1933, 901-908; Christoff, Phytopath. Zeitschr., 7, 1934, 521-536; S, 1935, 285-296; Thomas, Hilgardia, 10, 1937, 581-588. 48. Marmor fragariae H. {loc. cit., 78). From New Latin Fragaria, generic name of strawberry, from Latin fraga, straw- berries. Common name: Strawberry-crinkle virus. Hosts : ROSACEAE — Fragaria hy- brids, cultivated strawberries. Experi- mentally, also Fragaria vesca L., wood- land strawberry. Geographical distribution: United States, England. Induced disease: In cultivated straw- berry, crinkling and chlorosis of leaves. At first, minute chlorotic flecks appear in young leaves. These fiecks enlarge, and small necrotic spots may appear in their centers. Vein-clearing appears fre- quently. Affected foliage lighter and less uniformly green than normal. The vari- ety Royal Sovereign may appear normal through carrying this virus. Transmission : By aphid, Myzus fragae- folii Ckll. (= Capitophoriis fragariae Theob.) (APHIDIDAE). By grafting. Not by inoculation of expressed juice. Literature: Harris, Ann. Rept. East Mailing Res. Sta. for 1936, 1937, 201-211, 212-221; ibid., for 1937, 1938, 201-202; Harris and Hildebrand, Canad. Jour. Res., C, 15, 1937, 252-280; Ogilvie et al., Ann. Rept. Long Ashton Res. Sta. for 1933, 1934,. 96-97; Vaughan, Phytopath., 23, 1933, 738-740; Zeller, Oregon Agr. Exp. Sta., Sta. Bull. 319, 1933; Zeller and \\aughan, Phytopath., 22, 1932, 709-713. 49. Marmor marginans H. (loc. cit., 79). From Latin marginare, to provide with a margin. Common name: Strawberry j^ellow- edge virus. Hosts : ROSACEAE— Fragaria hy- brids, strawberries; Fragaria calif arnica C. and S. ; F . chiloensis Duch. (symptom- less). Experimentally, also Fragaria vesca L. ; F. virginiana Duch. (some clones appear to be immune to infection by runner inarching). Geographical distribution: United States, England, France, New Zealand. Induced disease : In strawberry, plant appears flat with outer zone of leaves more or less normal, central leaves dwarfed, yellow-edged, deficient in red pigmenta- tion. The variety Premier may carry this virus without showing any obvious manifestation of disease. Transmission : By aphid, Myzus fragae- folii Ckll. {APHIDIDAE). By graft- ing. Not b}' inoculation of expressed juice. Not through seeds from diseased plants. Literature : Chamberlain, New Zealand Jour. Agr., J^9, 1934, 226-231; Harris, Jour. Pom. and Hort. Science, 11, 1933, 56-76; Harris and Hildebrand, Canad. Jour. Res., C, 15, 1937, 252-280; Hilde- brand, ibid., C, 19, 1941, 225-233; Plaki- das, Phytopath., 16, 1926, 423-426; Jour. Agr. Res., 35, 1927, 1057-1090. 50. Marmor rubi H. (Holmes, loc. cit., 80; Poecile rubi McKinney, Jour. Wash- ington Acad. Science, 3J^, 1944, 148.) From Latin rubus, bramble bush. Common name : Red-raspberry mosaic virus. Hosts : ROSACEAE — Rubus idaeus L., red raspberry; R. occidentalis L., black raspberry. Geographical distribution: United States. Induced disease : In red raspberry, sys- temic chlorotic mottling, masked at high temperatures of summer. Foliage de- velopment delayed in spring. In some varieties, leaf petioles and cane tips die, 1196 MANUAL OF DETERMINATIVE BACTERIOLOGY canes remain short and become rosetted. Transmission: By aphids, principally Amphorophora rubi Kalt., but also A. rubicola Oestl. and A. sensoriaia Mason (APHIDIDAE). Not by inoculation of expressed juice. Literature: Bennett, Michigan Agr. Exp. Sta., Techn. Bull. 80, 1927; 125, 1932; Cooley, New York Agr. Exp. Sta. (Geneva), Bull. 675, 1936; Harris, .Jour. Pom. and Hort. Science, 11, 1933, 237-255 ; 17, 1940, 318-343; Rankin, New York Agr. Exp. Sta., Geneva, Bull. 543, 1927; New York Agr. Exp. Sta., Geneva, Tech. Bull. 175, 1931. 51. Marmor persicae H. (Holmes, loc. cit., 81 ; Flavimacula persicae McKinncy, Jour. Washington Acad. Science, 34, 1944, 149.) From New Latin Persica, former generic name of peach. Common name : Peach-mosaic virus. Hosts: ROSACEAE—Prunus persica (L.) Batsch, peach and nectarine, all tested varieties. Experimentally, also P. armeniaca L., apricot; P. communis Fritsch, almond; P. domestica L., plum and prune. Insusceptible species : Attempts to in- fect sweet and sour cherries have thus far failed. Geographical distribution : United States (Colorado, California, Utah, Ok- lahoma, Texas, New Mexico, Arizona). Induced disease : In peach, short inter- nodes in spring growth, sometimes break- ing in flower pattern, chlorotic mottling and distortion of foliage early in season, masking of leaf symptoms or excision of affected areas of leaf lamina in midsum- mer; fruit small, irregular in shape, un- salable. Some peach varieties are less damaged than others, but all are thought to be equally susceptible to infection, and equally important as reservoirs of virus when infected. In almond, experimen- tally, symptomless infections ; symptoms appear in some apricot and plum varieties when experimentally infected, not in others. Transmission : By budding and other methods of grafting. Not by inoculation of expressed juice. Not through soil. No insect vector is known. Not through pollen or seed from diseased plants. Thermal inactivation : Not demon- strated; virus not inactivated by tem- peratures effective in inactivating peach- 3'ellows virus. Literature : Bodine, Colorado Agr. Exp. Sta., Bull. 421, 1936; Bodine and Durrell, Phytopath., 31, 1941, 322-333; Cation, ibid., 2A, 1934, 1380-1.381; Christoff, Phytopath. Ztschr., 11, 1938, 360-422; Cochran, California Cultivator, 87, 1940, 164-165; Cochran and Hutchins, Phyto- path., 28, 1938, 890-892; Hutchins, Sci- ence, 76, 1932, 123; Hutchins et al., U. S. Dept. Agr., Circ. 427, 1937, 48 pp.; Kunkel, Am. Jour. Bot., 23, 1936, 683- 686; Phytopath., 28, 1938, 491-497; Thomas and Rawlins, Hilgardia, 12, 1939, 623-644; Valleau, Kentucky Agr. Exp. Sta., Bull. 327, 1932, 89-103. 52. Marmor astri H. {loc. cit., 83). Fi'om Latin astruin, star. Common name : Peach asteroid-spot virus. Host : ROSACE AE — Pruniis persica (L.) Batsch, peach. Geographical distribution : California. Induced disease : In peach, discrete chlorotic lesions spreading along veins, forming star-like spots ; developing leaves normal in appearance, becoming affected as they mature. Some chlorophyll re- tained in lesions as leaves turn yellow. Affected leaves shed early. Transmission: By grafting. Not by in- oculation of expressed juice. No insect vector is known. Literature : Cochran and Smith, Phyto- path., 28, 1938, 278-281. 53. Marmor rubiginosum Reeves. (Phytopath., 30, 1940, 789.) From Latin rubiginosus, rusty. Conunon name : Cherry rusty-mottle virus. Host : ROSACE AE — Primus avium L., sweet cherry. FAMILY MARMOEACEAE 1197 Geographical distribution : United States (Washington). Induced disease : In sweet cherry, chlorotic mottling 4 to 5 weeks after full bloom, first on small basal leaves, later on all leaves. The older affected leaves de- velop autumnal colors and absciss, 30 to 70 per cent of the foliage being lost. The remaining foliage appears somewhat wilted, shows increased mottling, chlo- rotic spots, and ai'eas becoming yellowisli brown, appearing rusty. Blossoms nor- mal. Fruits smaller than normal, in- sipid, not misshapen. Growth rate of tree reduced slightly. Transmission: By grafting. Not by inoculation of e.xpressed juice. No insect vector is known. Literature : Reeves, Phytopath., 30, 1940, 789 (Abst.); Jour. Agr. Res., 62, 1941, 555-572 (see 566-567). 54. Marmor cerasi Zeller and Evans. (Phytopath., 31, 1941, 467.) From Latin cerasus, cherry tree ; originally spelled cerasae, by error. Common name : Cherry mottle-leaf virus. Hosts : ROSACEAE — Primus avium L., sweet cherry; P. emarginata (Dougl.) Walp., wild cherry. Experimentally, also P. cerasus L. (tolerant) and P. ma- haleb L. (tolerant). Geographical distribution : United States (Washington, Oregon, Idaho, Cali- fornia) and Canada (British Columbia). Induced disease : In sweet cherry, chlo- rotic mottling ; leaves puckered, wrinkled, distorted, not perforated. Blossoms not affected. Fruit small, hard, insipid, un- even or delayed in ripening. Crop re- duced. Branches shortened, tree even- tually stunted. Transmission : By budding. No insect vector is known. Not by the black cherry aphid, Mysws cerasi (F.) (APHI- DIDAE). Not by inoculation of ex- pressed juice. Thermal inactivation : Not demon- strated ; not at 46° C in 60 minutes nor at 49° C in 10 minutes in bud sticks. Literature : Reeves, Washington State Hort. Assoc. Proc, 31, 1935, 85-89; Jour. Agr. Res., 62, 1941, 555-572; Zeller, Ore- gon State Hort. Soc. Report, 26, 1934, 92-95; Phytopath., 31, 1941, 463-467. 55. Marmor lineopictum Cation (Phytopath., 81, 1941, 1009.) From Latin linea, line, and pictus, ornamented. Common names : Prunus line-pattern virus, peach line-pattern virus. Hosts : ROSACEAE — Prunus salicina Lindl., Japanese plum; P. mahaleb L., Mahaleb chei'ry; P. persica (L.) Batsch, peach (= Amy gdalus persica \j.). Geographical distribution : United States (Kentucky, Michigan, California, Ohio; perhaps widely distributed). Induced disease : In peach and Ma- haleb cherry, light-colored line patterns or faint chlorotic mottling, tending to become masked as leaf becomes old. In peach, affected foliage sometimes less glossy than normal. In Prunus salicina, no disease manifestations usually ; rarely, chlorotic mottling on a few leaves. Transmission : By grafting. No insect vector is known. Literature : Berkeley, Div. of Bot. and Plant Path., Science Service, Dominion Dept. Agr., Ottawa, Canada, Publ. 679, 1941; Cation, Phytopath., 31, 1941, 1004- 1010; Thomas and Rawlins, Hilgardia, 12, 1939, 623-644 ; Valleau, Kentucky Agr. Exp. Sta., Res. Bull. 327, 1932, 89-103. 56. Marmor pallidolimbatus Zeller and Milbrath. (In Handbook of Virus Dis- eases of Stone Fruits in North America, Michigan Agr. Exp. Sta., Miscell. Publ., May, 1942, 50; Phytopath., 32, 1942, 635.) From Latin pallidus, pale, and limbatus, bordered. Common name : Cherr}^ banded-chloro- sis virus. Hosts: ROSACEAE — Prunus serru- lata Lindl., flowering cherrj^; P. avium L., Mazzard cherry. Geographical distribution : United States (Pacific Northwest). Induced disease : In flowering cherry. 1198 MANUAL OF DETERMINATIVE BACTERIOLOGY chlorotic bands surrounding discolored areas on leaves. In Mazzard cherry, dwarfing of whole plant, chlorotic bands on leaves. Transmission : By budding, even in the absence of survival of inserted buds. 57. Marmor nerviclarens Zeller and Evans. (Phytopath., 31, 1941, 467.) From Latin nervus, sinew or nerve, and clarere, to shine. Conamon name : Cherry vein -clearing virus. Hosts : ROSACE AE — Primus avium L., sweet cherry. Perhaps also P. serru- lata Lindl. and P. domestica L., on which symptoms similar to those induced by this virus have been observed. Geographical distribution : United States (Oregon, Washington). Induced disease : In sweet cherry, clearing of veins throughout each leaf or only in localized areas. Margins of leaves irregular, most indented where clearing of veins is most conspicuous. Elongated, elliptic, or slot-like perforations occur in some leaves. Affected leaves usually narrow. Enations occur as small blis- tered proliferations on lower side of main veins. Upper leaf surface silvery by re- flected light. By midsummer, leaves droop and appear somewhat wilted ; they may fold along the midvein. Internodes short; increased number of buds, spurs, or short branches at nodes ; resetting more pronounced on some branches than on others, mostly at end of year-old wood. In advanced disease, fruits pointed, small, flattened on suture side with swol- len ridge along suture. Blossoms ab- normally abundant, crop of fruit reduced or wanting. Transmission : By grafting. Not dem- onstrated by inoculation of expressed juice. No insect vector is known. 58. Marmor viticola H. (loc. cit., 83). From Latin vitis, vine, and -cola, in- habitant of. Common name : Vine -mosaic virus. Host : VITACEAE—Viiis vinifera L., grape . Geographical distribution : France, Italy, Bulgaria, Czechoslovakia. Induced disease : In grape, various modifications of systemic chlorotic mot- tling, and red pigmentation of parts of leaves with subsequent drying and drop- ping out of affected spots. Leaves de- formed, crimped between main veins. Growth restricted. Transmission : By inoculation of ex- pressed juice and by pruning. Literature: Blattny, Vinafsky obzor., 25, 1931, 4-5 (Cent. f. Bakt., II Abt., 84, 1931, 464); Ochrana Rostlin, 13, 1933, 104-105 (Rev. Appl. Mycol., 13, 1934, 421) ; Gigante, Boll. Staz. Pat. Veg. Roma, n. s. 17, 1937, 169-192 (Rev. Appl. Mycol., 17, 1938, 221) ; Pantanelli, Malpighia, 24, 1911, 497-523; ^5, 1912, 17-46; Stranak, II Congr. Intern. Path. Comp. Paris, 1931, 367-378; Ochrana Rostlin, 11, 1931, 89-98 (Rev. Appl. Mycol., 11, 1932, 280) ; Vielwerth, Ochrana Rostlin, 13, 1933, 83-90 (Rev. Appl. Mycol., 13, 1934, 421-422). 59. Marmor santali H. {loc. cit., 94). From New Latin, Santalum, generic name of sandal. Common name : Sandal leaf -curl virus. Host : S AN TAL AC EAE— Santalum album L., sandal. Geographical distribution : India. Induced disease: In sandal, leaves small, curled, wrinkled, thickened, brit- tle, abscissing. Systemic chlorotic mot- tling. Internode length normal. In- fected twigs produce both flowers and fruits. Transmission : By ring bark-grafts. Not by inoculation of expressed juice. No insect vector is known. Literature : Venkata Rao, Mysore San- dal Spike Invest. Comm., Bull. 3, 1933. 60. Marmor secretum Bennett. (Phy- topath., 34, 1944, 88). From Latin se- cretus, hidden. Common name : Dodder latent-mosaic virus. Hosts: CONVOLVULACEAE—Cus- cuta californica Choisy, dodder. Experi- FAMILY MARMORACEAE 1199 mentally, also CHENOPODIACEAE— Beta vulgaris L., sugar beet ; Chenopo- dium albvm L., lamb's quarters; C. murale L., sowbane. CONVOLVULA- CEAE — Citsciita campesiris Yuncker; C. subinclusa Dur. and Hilg. CRUCI- FERAE — Brassica incana (L.) F. W. Schultz, mustard (tolerant). CUCUR- BIT ACE AE — Cucumis melo L., canta- loupe. P HYTOLACC ACE AE— Phyto- lacca americana L., pokeweed. PLAN- T AGIN ACE AE—Plantago major L., plantain. POL YGON ACEAE—Fago- pyru7n escidentmn Moench, buckwheat ; Polygonum pennsylvanicinn L., knot- weed. PRIM ULACEAE—Samolus florihundus HBK., water pimpernel. SOLA NA CEAE — Lycopersicon esculen- tum Mill., tomato ; Nicotiana glauca Gra- ham (tolerant); N. palmeri Gray; A''. rustica L. (tolerant); A'', tabacum L. (tolerant); Solanum tuberosum L., po- tato. UMBELLIFERAE—Apium graveolens L., celery. Insusceptible species : COMPOS I TAE — Helianthus annuus L., sunflower; Lac- tuca sativa L., lettuce. CR UC I FERAE — Brassica oleracea L. , cabbage . POL Y- GONACEAE — Eriogonum, fasciculatum Benth., California buckwheat. SCRO- PH ULARIACEAE—Verbascum thap- sus L., mullein. SOLAN ACE AE— Atropa belladonna L., belladonna. Geographical distribution : United States (California). Induced disease : In dodder, no symp- toms. In sugar beet, experimentally, temporary systemic chlorotic spotting; occasional faded areas in leaves in subse- quent chronic stage of disease. In canta- loupe, experimentally, chlorotic spotting, reduction in leaf size, death of some leaves, stunting of plant; melons small and of poor . quality. In celery, experi- mentally, systemic chlorosis followed by dwarfing and mottling with subsequent apparent recovery. Transmission : By dodder, Cuscuta cali- fornica, C. campestris, and C. subinclusa. By inoculation of extracted juice to some, but not to other, host plants ; Phytolacca americana is readily infected by rubbing methods in the presence of a small amount of abrasive, and develops numerous necrotic primary lesions that serve for quantitative estimation of concentration of virus in inoculum. Through seeds from infected plants of dodder, Cuscuta campestris; not through seeds from dis- eased cantaloupe, buckwheat, or poke- weed plants. No insect vector is known. Thermal inactivation : At 56 to 60° C in 10 minutes. Filterability : Passes celite and Berke- feld N and W filters. Other properties : Infective in dilu- tions to 1 : 3000. Inactivated by drying and by storage in expressed pokeweed juice, within 48 hours. 61. Marmor pelargonii spec. nov. From New Latin Pelargonium, generic name of common geranium. Common names : Pelargonium leaf -curl virus ; virus of dropsy or Krauselkrank- heit of geranium. Host : GERAN I ACE AE— Pelargon- ium hortorum Bailey, geranium. Induced disease : In geranium, circular or irregular chlorotic spots, sometimes stellate or dendritic, | to 5 mm in di- ameter, centers becoming brown with chlorotic border ; severely affected leaves become yellow and drop; spotted leaves ruffled, crinkled, malformed, small, some- times puckered and splitting. Petioles and stems show corky, raised, necrotic streaks; tops may die. Disease most severe in spring, inconspicuous in sum- mer. Transmission : By grafting. Not by inoculation of expressed juice nor by use of knife to prepare cuttings for propaga- tion. Not through seed. No insect vec- tor is known. Literature : Berkeley, Canad. Hort. and Home Mag., 1938, 1938, 1-4; Blattn^, Ochrana Rostlin, 13, 1933, 145 (Rev. Appl. Mycol., 13, 1934, 378-379) ; Bremer, Blumen-u. Pflanzenbau, J^8, 1933, 32-33 (Rev. Appl. Mycol., 12, 1933, 514); Halstead, New Jersey Agr. Exp. Sta., Rept. 14, 1893, 432-433; Jones, Washing- ton Agr. Exp. Sta., Bull. 390, 1940; 1200 MANUAL OF DETERMINATIVE BACTERIOLOGY Laubert, Gartenwelt, 31, 1927, 391 ; Pape, ibid., 26, 1927, 329-331 ; 32, 1928, 116-117; Pethybridge and Smith, Gard. Chron., 92, 1932, 378-379; Schmidt, Gartenwelt, 81, 1932, 40; Seeliger, Nachrichtenbl. Deutsch. Pflanzenschutzdienst, 6, 1926, 63-64; Tuimann, Gartenwelt, 31, 1927, 375-376; Verplancke, Bui. 01. Sci. Acad. Roy. de Belgique, Ser. 5, 18, 1932, 269- 281 (Rev. Appl. Mycol., 11, 1932, 649- 650). 62. Marmor angliae H. {loc. cit., 48). From Latin Anglia, England. Common name : Potato-paracrinkle virus. Hosts: SOLA N ACE AE—Solanum tu- berosum L., potato. Experimentally, also Datura stramonium L., .Timson weed. Insusceptible species: SOLAN A- CEAE — Nicotiana iabacum L., tobacco. Geographical distribution : England. Induced disease : In potato, masked in all plants of the variety King Edward. Chlorotic mottling with some necrosis in the varieties Arran Victory and Arran Chief. Chlorotic mottling only in Arran Comrade, Majestic, and Great Scot pota- toes. Two varieties, Sharpe's Express and Epicure, are said to be resistant. Transmission: By grafts. Not by in- oculation of expressed juice. No insect vector is known. Literature: Dykstra, Phytopath., 26, 1936, 597-606; Salaman and Le Pelley, Proc. Roy. Soc. London, Ser. B, 106, 1930, 140-175. 63. Marmor aevi spec. nov. From Latin aevum, old age, in reference to the obvious involvement of old, but not of young, delphinium leaves. Common name : Celery-calico virus. Hosts: CUCURBIT ACE AE—Ciict(- mis sativus L., cucumber; C. melo L., cantaloupe; Cucurbita pepo L., summer crookneck squash. RANUNCULA- CEAE — Delphinium chinensis ; D.formo- sum, hardj" larkspur; D. grandiflorum; D. parryi; D. zaliL SOLAN ACE AE—Ly- copersicon esculentum. Mill., tomato. UM BELLI FERAE— Apium graveolens L., celery. Experimentally, also SOLA- NACEAE — Nicotiana tabacum L., to- bacco; Petunia hybrida Vilm., petunia. V lOL ACE AE— Viola cornuta L. Geographical distribution : United States. Induced disease : In celery, clearing of veins, puckering and downward cupping of younger leaves, green islands of tissue in lemon-yellow areas of outer leaves, green and yellow zigzag bands on leaflets. In delphinium, basal and middle leaves with pale-orange, amber, or lemon-yellow areas; younger leaves normal green; chlorotic ring and line patterns. Transmission : By inoculation of ex- pressed juice in the presence of finely jiowdered carborundum. By aphids : Aphis apigraveolcns Essig, celery leaf aphid; A. apii Theob., celery aphid; A. ferruginea-striala Essig, rusty-banded aphid; A. gossypii Glov., cotton aphid; A. middletonii Thomas, erigeron root aphid; Myzus circumflexus (Buckt.), lily aphid; M. convolvidi (Kalt.), foxglove aphid; M. persicae (Sulz.), green peach aphid; Rhopalosiphum melliferum (Hot- les), lioneysucklc aphid (APHIDI- DAE). Literature : Severin, Hilgardia, IJ^, 1942, 441-464 ; Severin and Freitag, Phy- topath., 25, 1935, 891 (Abst.); Hilgardia, n, 1938, 493-558. 64. Marmor raphani spec. nov. From Latin raphanus, radish. Common name : Radish-mosaic virus. Hosts : CR UC I FERAE— Raphanus sativus L., radish. Experimentally, also CRUCIFERAE—Brassica oleracea L.; B. nigra (L.) Koch; B. alba (L.) Boiss; B. arvensis (L.) Ktze. ; B. pe-tsai Bailey; B. juncea (L.) Coss; B. rapa L.; B. ad- pressa Boiss ; Capsclla bursa-pastoris (L.) Medic; Malcomia maritima R. Br.; M. bicornis DC. CHENOPODIACEAE— Chenopodium album L.; C. murale L. ; Spinacia oleracea L. RANUNCULA- CEAE — Delphinium ajacis L. SOLA- N ACE A E— Nicotiana glntinosa L. ; A'. FAMILY MARMORACEAE 1201 langsdorffii Weinm. ; .V. riistica L.; N. tahacum L. Geographical distribution : United States (California). Induced disease : In radish, systemic chlorotic spotting followed by chlorotic mottling of foliage; little or no leaf dis- tortion; plants not stunted. Transmission : Bj' inoculation of ex- pressed juice. No insect vector is known ; not by the cabbage aphid, Brevi- coryne brass icac (L.);the false cabbage aphid, Lipaphis pseudobrassicae (Davis) ; or the green peach aphid, Myzus persicac (Sulz.) (APHIDIDAE). Not through seeds from diseased radish plants. Thermal inactivation : At 65 to 68° C in 10 minutes. Literature : Tompkins, Jour. Agr. Res., 58, 1939, 119-130. 65. Marmor primulae spec. nov. From New Latin Primula, generic name of primrose. Common name : Primrose -mosaic virus. Hosts: PRIM UL ACE AE— Primula obconica Hance. Experimentally, also P. malacoides Franch. and P. sinensis Lindl. Insusceptible species: BEGONIA- CEAE — Begonia semper flor ens Link and Otto. BORAGINACEAE—Mijosotis alpestris Schmidt. CAMPANULA- CEAE — Campanula medium L. CARY- OPHYLLACEAE—Dianthus barbatus L. CHENOPODIACEAE—Spinacia oleracea L. COMPOSITAE—Bellis perennis L. ; Callistephus chinensis Nees ; Gerbera jamesonit Hook. ; Lactuca saliva L. ; Senecio cruentus DC; Tagetes patula L. CRUCIFERAE—Brassica oleracea L. ; B. pe-tsai Bailey ; B. rapa L. ; Matthi- ola incana R. Br.; Raphanus saiivus L. CUCURBI TA CEA E~Cucu m is saiiv us L.; Cucurbita pepo L. EUPHORBIA- CEAE — Ricinus communis L. GR AM- INE AE—Zea mays L. LEGUMINO- SAE — Pisum sativum. L.; Vicia faba L. ; Vigna sinensis (Torner) Savi. LO- BEL I ACE A E— Lobelia hybrida Hort. P AP AVER ACE AE—Papaver orientale L. PRIM UL ACE AE—Anagallis ar- vensis L. ; Cyclamen indicum L. ; Primula auricula L. ; P. veris L. RANUNCU- LACEAE — Anemone coronaria L. ; Del- phinium cultorum Voss; Ranunculus asiaticus L. RESEDACEAE—Reseda odorata L. ROSACEAE — Geum chiloense Balb. SCROPHULARI ACE AE— An- tirrhinum majus L. ; Pentstemon barbatus Nutt. SOLA N ACE AE— Capsicum, fru- tescens L. ; Datura stramonium L. ; Lyco- persicon esculentum Mill.; Nicotiana glutinosa L. ; N . tabacum L. ; Solanum tuberosum L. TROPAEOLACEAE— Tropaeolum majus L. UMBELLI- FERAE — Apium graveolens L. VER- BE N ACE A E— Verbena hybrida Voss. V lOL ACE AE— Viola tricolor L. Geographical distribution : United States (California). Induced disease : In Primula obconica, chlorosis, stunting, rugosity with upward, or occasionally downward, cupping of leaves. Petioles and peduncles short- ened; flowers reduced in size, broken in color (white-streaked). Leaves coarsely mottled with yellow-green, leaving green islands ; tips of leaves sometimes nar- rowed. Transmission : By inoculation of ex- pressed juice, in the presence of 600-mesh powdered carborundum. Not by aphids, Myzus persicae (Sulz.) and M. circum- flexus (Buckt.) {APHIDIDAE). No insect vector is known. Probably not through seeds. Thermal inactivation : At 50° C, not 48° C, in 10 minutes. Other properties : Infective after 24, not 48, hours in vitro. Infective after 1:10 dilution. Literature : Tompkins and Middleton, Jour. Agr. Res., 63, 1941, 671-679. 66. Marmor caricae (Condit and Home) comb. nov. {Ficivir caricae Condit and Home, Phytopath., 81, 1941, 563.) From Latin carica, a kind of dried fig. Common name : Fig-mosaic virus. Hosts : MORACEAE—Ficus carica L., 1202 MANUAL OF DETERMINATIVE BACTERIOLOGY fig; F. altissima Blume; F. Jcrishna; and F. tsiela Roxb. Geographical distribution : United States (California, Texas), England, Puerto Rico, China, New South Wales, Western Australia. Induced disease : In fig, systemic chlo- rotic spotting and mottling of foliage ; some severe leaf distortion. Fruits some- times afi"ected, bearing light circular areas, rusty spots, being deformed or dropped prematurely. Necrotic lesions on profichi of Samson caprifigs also have been attributed to action of this virus. Transmission : By budding. No insect vector is known; mites have been sus- pected as possible vectors. Literature: Condit and Home, Phyto- path., 23, 1933, 887-896; 31, 1941, 561- 563 ; 33, 1943, 719-723 ; Ho and Li, Lingnan Science Jour., 15, 1936, 69-70; Pittman, W. Austral. Dept. Agr. Jour., 12, 1935 196. 67. Marmor italicum (Fawcett) comh. nov. {Citrivir italicum Fawcett, Phyto- path., 31, 1941, 357.) Specific epithet meaning "pertaining to Italy." Common name : Citrus infectious-mot- tling virus. Host: RU T ACE AE— Citrus auran- tium L., sour orange. Geographical distribution : Italy. Induced disease : In sour orange, white, pale green, or yellow irregular areas in leaves, leaving narrow green bands along midrib; leaves blistered and distorted. Transmission : The aphid, Toxoptera aurantii (Phytopath., 2J^, 1934, 661), has . been suspected as vector. Literature: Fawcett, Phytopath., 31, 1941, 356-357; Petri, Bol. Staz. Pat. Veg. Roma, n. s. 11, 1931, 105-114. Note : Several additional species were described too late for complete systematic treatment here. They are plain's wheat mosaic virus, Marmor campestre McKinney (Jour. Washington Acad. Sci., SJj., 1944, 324) with varieties iypicum McKinney and galhinmn McKinney, re- spectively causing light-green mosaic and severe yellow mosaic of wheat in Kansas; wheat streak-mosaic virus, Mar- mor virgatum McKinney (ibid., 34, 1944, 324) with varieties typicuni McKinney and viride McKinney (ibid., 34, 1944, 325), respectively causing yellow streak- mosaic and green streak-mosaic of wheat in Kansas; Agropyron-mosaic virus, Marmor agropyri McKinney (ibid., 34, 1944, 326), with varieties typicum McKinney and flavum McKinney, re- spectively causing green-mosaic mottling and yellow-mosaic mottling in the grass Agropyrou repens (L.) Beauv. in Vir- ginia; also a virus, Flavimacula ipomeae Doolittle and Harter (Phytopath., 35, 1945, 703), causing feathery mottle of sweet potato in Maryland [see Marmor persicae for treatment of a virus that was assigned as type of Flavimacula McKinney (Jour. Washington Acad. Sci., 34, 1944, 149), a genus originally differ- entiated from Marmor as containing viruses not yet inoculable save by tissue union ; a natural group of viruses may be represented but their characteristics and affiliations seem not yet clear]. Genus II. Acrogenus Holmes, {hoc. cit., 110.) Viruses of the Spindle-Tuber Group, inducing diseases characterized by abnormal growth habit of host plants without chlorotic or necrotic spotting, systemic chlorosis, witches'-broom formation, or production of galls. Generic name from Greek, meaning point- or peak-producing, in reference to shape of potatoes affected by potato spindle- tuber virus. The type species is Acrogenus solani Holmes. FAMILY MARMORACEAE 1203 Key to the species of genus Acrogenus I. Infecting potato. II. Infecting black currant. 1. Acrogenus solani Holmes. (Handb. Phytopath. Viruses, 1939, HI.) From New Latin Solanum, generic name of potato. Common names : Potato spindle-tuber virus, potato spindling-tuber virus, po- tato marginal leaf -roll virus. Host: SOLAN AC EAE— Solanum tu- berosum L., potato. Geographical distribution : United States and Canada. Induced disease : Plants erect, stiff, spindly, lacking vigor. Leaves small, erect, darker green than normal. Peti- oles sometimes slender, brittle. Tubers long, cylindrical, irregular in shape, tap- ered at ends, smooth and tender-skinned, of softer than normal flesh in spring. Eyes of tuber conspicuous. Transmission: By inoculation of ex- pressed juice; by use of contaminated knife in cutting successive tubers before planting; by contacts of freshly cut seed pieces. By aphids, Myzus persicae (Sulz.) and Macrosiphum solanifolii Ashm.i=M.geiKoch) (APHIDIDAE). Also by certain leaf-eating insects. Thermal inactivation : At 60 to 65° C in 10 minutes (in tuber tissues). Literature : Bald et al., Phytopath., 31, 1941, 181-186; Folsom, Maine Agr. Exp. Sta., Orono, Bull. 312, 1923; Goss, Phyto- path., 16, 1926, 233, 299-303; 18, 1928, 445-448; Nebraska Agr. Exp. Sta., Res. Bull. 47, 1930; 53, 1931; Jaczewski, La Defense des Plantes, Leningrad, 4, 1927, 62-77 (Rev. Appl. Mycol., 6, 1927, 572- 573, Abst.) ; McLeod, Canad. Exp. Farms, Div. Bot., Rpt. for 1926, 1927. 1. Acrogenus solani. 2. Acrogenus ribis. Strains : A strain causing unmottled curly dwarf of potato has been given a varietal name to distinguish it from the type, var. vulgaris H. (loc. cit.. Ill) : la. Acrogenus solani var. severus H. (loc. cit., 112). Inducing symptoms in potato on the whole more severe than those caused by the type strain. Common name : Unmottled curly-dwarf strain of potato spindle -tuber virus. (Goss, Nebraska Agr. Exp. Sta., Res. Bull. 47, 1930; 53, 1931 ; Schultz and Fol- som, Jour. Agr. Res., 25, 1923, 43-118.) 2. Acrogenus ribis H. (loc. cit., 112). From Latin ribes, currant. Common name : Black-currant rever- sion-disease virus. Host: SAXIFRAGACEAE—Ribes nigrum L., European black currant. Geographical distribution : British Isles. Induced disease : In European black currant, leaves abnormally narrow and flat, small veins few. Flowers sometimes nearly transparent, smooth, sepals brightly colored beneath. Flowers and small fruits fall. Stems less woody than normal, with tendency to excessive gum production. Transmission : By grafting. By big- bud mite, Eriophyes ribis (ERIOPHYI- DAE). Not by inoculation of expressed juice. Not through soil. Not through seeds from diseased plants. Literature : Amos and Hatton, Jour. Pom. and Hort. Science, 6, 1926, 167-183; Amos et al., in East Mailing Res. Sta., 15th Ann. Rpt., 1928, 43-46; Lee, Ann. Appl. Biol., 9, 1935, 49-68. Genus III. Cerium Holmes. {Loc. cit., 119.) Viruses of the Leaf -Roll Group, inducing diseases usually characterized by thicken- 1204 MANUAL OF DETERMINATIVE BACTERIOLOGY ing and rolling of leaves. Foliage leathery. Sometimes conspicuous phloem necrosis. Generic name from Latin coriuvi, leather. The type species is Corium solani Holmes. Key to the species of genus Corium. I. Infecting potato. II. Infecting beet. III. Infecting raspberry. 1. Corium solani Holmes. (Handb. Phytupath. Viruses, 1939, 120.) From New Latin Solanum, generic name of potato. Common name: Potato leaf -roll virus. Hosts: SOLAN AC EAESolanum tu- berosum L., potato. Experimentally, also other solanaceous species. Datura stramonium L., Jimson weed; Lycopersi- con esculentum Mill., tomato; Solanun) dulcamara L., bittersweet; S. villosum. Insusceptible species : CHENOPODI- ACEAE — Beta vulgaris L., beet. Geographical distribution: North America, France, British Isles; probably wherever potatoes are grown. Induced disease : In potato, leaves thick, rigid, leather}^, and rolled, their starch content excessive. Plants dwarfed. Tubers few, small, crisp. Tubers of some varieties show conspicuous phloem necrosis, germinate with spindling sprouts. Transmission: By aphid, Myzus persi- cae (Sulz.) {APIilDIDAE), with incu- bation period of 24 to 48 hours. Also by Myzus convolvuli (Kalt.) (= M. pseudo- solaniT\i(ioh.),M. circumflexus (Buckt.), Macrosiphum solanifolii Ashm., and Aphis abbreviata Patch (APHIDI- DAE). By grafting. Not by inocula- tion of expressed juice. Literature : Artschwager, Jour. Agr. Res., 15, 1918, 559-570; 2i, 1923, 237-245; Dykstra, ibid., 47, 1933, 17-32; Elze, Phytopath., 21, 1931, 675-686; Folsom, Maine Agr. Exp. Sta., Bull. 297, 1921, 1. Corium solani. 2. Corium betae. 3. Corium rubi. 4. Corium ruborurn. 37-52 ; 410, 1942, 215-250 ; Murphy, Scient . Proc. Roy. Dublin Soc, 17, 1923, 163- 184; Murphy and M'Kay, ibid., 19, 1929, 341-353 ; Schultz and Folsom, Jour. Agr. Res., 21, 1921, 47-80; Smith, Ann. Appl. Biol., 16, 1929, 209-229; /8, 1931, 141-157; Stevenson et al.. Am. Potato Jour., 20, 1943, 1-in. 2. Corium betae spec. nov. From Latin beta, beet. Common names : Sugar-beet yellows virus, beet-yellows virus, jaunisse virus, vergelingsziekte virus. Hosts: CHENOPODI ACE AE— Beta vulgaris L., beet; B. maritima L. ; B. cicla; Atriplex hortensis L. ; A. sibirica L. ; Clicnopodium album L., lamb's quar- ters; Spinacia oleracea L., spinach. AMARANTHACEAE —Amaranthus retroflexus L. Insusceptible species: SOLAN A- CEAE — Solanum tuberosum L., potato; all other tested solanaceous species. Geographical distribution : Belgium, Netherlands, Denmark, England; per- haps Germany and the United States. Induced disease : In beet, young leaves little affected; older leaves yellow, brit- tle, short, thick, containing excessive amounts of carbohydrates; necrosis in secondary phloem. In spinach, yellow- ing, necrosis between veins on old leaves. Transmission : Not by inoculation of expressed juice. By aphids, Myzus per- sicae (Sulz.), Aphis fabae Scop., Macro- siphum solanifolii Ashm., and Ardacor- FAMILY MARMORACEAE 1205 thurn solani Kalt. (APHIDIDAE) ; virus is not transmitted by these aphids to their descendants. Not through seeds of beet. Virus overwinters in beets stored for subsequent use in seed produc- tion. Serological relationships: Specific pre- cipitating antiserum effective with crude sap of diseased, not healthy, plants and with sap of diseased plants after pas- sage through a Chamberland Li, not Ls, filter candle; ineffective with sap from beet plants suffering from mosaic. Thermal inaetivation: Virus heated to about 52°C no longer precipitates with specific antiserum. Literature: Kleczkowski and Watson, Ann. Appl. Biol., 31, 1944, 116-120; Peth- erbridge and Stirrup, London, Ministry Agr. and Fisheries, Bull. 93, 1935; Quan- jer and Roland, Tijdschr. Plantenziek- ten, 42, 1936, 45-70; Roland, ibid., 45, 1939, 1-22, 181-203; Schreven, Meded. Inst, voor Suikerbietenteelt, Bergen op Zoom, 6, 1936, 1; Watson, Proc. Roy. Soc. London, Ser. B, 128, 1940, 535-552; Ann. Appl. Biol., 29, 1942, 358-365. 3. Corium rubi H. {loc. cit., 121). From New Latin Rubus, generic name of raspberry, from Latin rubus, bramble bush. Common name : Raspberry leaf -curl virus. Host : ROSACE AE— Rubus idaeus L., red raspberry. Insusceptible species : ROSACE AE — Rubus occidentalis L., black raspberry; R. neglectus Peck, purple raspberry. Geographical distribution : Unitetl States, not in England. Induced disease : In red raspberry, veins retarded in growth, causing down- ward curling of leaf margins and crinkling of leaf lamina. Foliage dark green, dry in appearance, not wilting readily. In late summer, leaves bronzed, leaf surface glis- tening. Diseased canes easily winter- killed. Berries small and poor. The English variety Lloyd George is intol- erant of the disease and is killed. Transmission : By aphid. Aphis rubi- cola Oestl. (== A. rubiphila Patch) (APHIDIDAE). Not by inoculation of expressed juice. Literature : Bennett, Michigan Agr. Exp. Sta., Tech. Bull. 80, 1927; Phyto- path., 20, 1930, 787-802, Harris, East Mailing Res. Sta., Ann. Rpt. for 1934, 1935; Rankin, New York Agr. Exp. Sta., Geneva, Tech. Bull. 175, 1931. Strains : A strain differing from the type, var. alpha H. {loc. cit., 121), has been given a varietal name derived frona its common name, raspberry beta-curl virus : 3a. Corium rubi var. beta H. {loc. cit., 122). Infecting black and purple rasp- berries, as well as the red raspberry, which alone is susceptible to the type strain, raspberry alpha-curl virus. (Ben- nett, Phytopath., 20, 1930, 787-802.) 4. Corium ruborum (Zeller and Braun) comb. nov. (Minuor ruborum Zeller and Braun, Phytopath., 33, 1943, 161.) From Latin rubus, bramble bush. Common name : Raspberry decline-dis- ease virus. Host : ROSACE AE — Rubus idaeus L., red raspberry. Geographical distribution : United States (Oregon). Induced disease : In Cuthbert rasp- berry, shoots retarded in spring, reddish; leaves in autumn rolled downward, fluted along veins, less green than normal be- tween veins, slightly bronzed along mar- gins and crests between veins. Inter- nodes shortened near tips of canes. Af- fected canes small, weak, not hardy in winter. Small roots and feeder rootlets fewer than in healthy plants. Disease progressive over about three years. Fruits small, irregular, tending to be globose, crumblj^ when ripe, worthless. Transmission: By grafting. No insect vector is known. 1206 MANUAL OF DETERMINATIVE BACTERIOLOGY Genus IV. Nanus Holmes, {hoc. cil., 123.) Viruses of the Dwarf-Disease Group, inducing diseases characterized by dwarfing of host plants or by growth of adventitious shoots with short internodes; chlorotic mottling absent. Generic name from Latin nanus, dwarf. The type species is Nanus loganobacci Holmes. Key to the species of genus Nanus. I. Infecting rosaceous plants. A. In loganberry and Phenomenal berry. 1. Nanus loganobacci. B. In black raspberry. C. In peach. D. In ocean spray. E. In strawberry. F. In prune and plum. II. Infecting graminaceous plants. A. In sugar cane. 1. Nanus loganobacci Holmes. (Handb. Phytopath. Viruses, 1939, 124.) From New Latin loganobaccus, specific epithet of loganberry, Riibus loganobaccus Bailey. Common name: Loganberry -dwarf virus. Hosts: ROSACEAE—Rubus logano- baccus Bailey, loganberry and Phenom- enal berry. Geographical distribution: United States (Oregon, Washington, and Cali- fornia). Induced disease : In Phenomenal berry, leaves small, obovate, rigid, new canes short, spindly. In young plants, some necrosis along and between veins, leaves crinkled, finer veins chlorotic. Stems not streaked or mottled, normal in color. In late stages, canes very short, inter- nodes short. Sepals and petals of flowers small. Fruit of fair size, but druplets ripen unevenly and tend to fall apart when picked. Loganberry is less sus- ceptible than the Phenomenal berry but is similarly affected. 2. Nanus orientalis. 3. Nanus mirabilis. 4. Nanus holodisci. 5. Nanus fragariae. 6. Namis cupitliformans. 7. Nanus prrinil 8. Nanus sacchari. Transmission : By aphid, Capitophorus tetrahodus (APHIDIDAE). Not by inoculation of expressed juice. Literature: Zeller, Phytopath., 15, 1925, 732 (Abst.) ; 17, 1927, 629-648. 2. Nanus orientalis H. {loc. cit., 124). From Latin orientalis, eastern. Common names : Raspberry -streak vi- rus, raspberr}'^ eastern blue-stem virus, raspberry rosette virus. Host: ROSACEAE—Rubus occiden- talis L., black raspberry. Insusceptible species : ROSACEAE — Rubus idaeus L., red raspberry; R. pho- enicolasius Maxim., Japanese wineberry. Geographical distribution : United States. Induced disease : In black raspberry, plants stunted, becoming smaller in suc- cessive seasons; leaves usually curled, close together on canes, dark green, often twisted so as to be upside down. New canes show bluish violet dots, spots, or stripes near their bases and sometimes also on branches or on fruiting spurs. FAMILY MARMORACEAE 1207 Fruit inferior in size, quality, and quan- tity. Plants live only 2 or 3 years after infection on the average. Strains: A strain of this virus is be- lieved responsible for mild streak of black raspberries, in which purple to violet, greenish brown, or bluish streaks on canes are narrowly linear or elliptical in form and often very faint; when the bloom is rubbed off, the lesions appear as though watersoaked and discolored. Leaves are slightly curled, their veins cleared. Fruits are dry and dull in lustre, even while still red, and of poor flavor when ripe. Literature: Bennett, Michigan Agr. Exp. Sta., Tech. Bull. 80, 1927; Wilcox U. S. Dept. Agr., Dept. Circ. 227, 1923; Woods and Haut, U. S. Dept. Agr., Plant Dis. Rpt., ?4, 1940, 338-340. 3. Nanus mirabilis H. {loc. cit., 126). From Latin mirabilis, strange. Conamon name : Peach phony-disease virus. Hosts : ROSACE AE — Primus persica (L.) Batsch, peach. Experimentally, also other Prunus species. Geographical distribution : United States (Georgia, Alabama, Florida; sparsely also in Mississippi, Tennessee, South Carolina, Louisiana, Texas, Ar- kansas, Missouri). Induced disease : In peach, tree dwarfed, foliage abnormally green, fruit small ; flecks in wood, especially in roots ; sections of roots show characteristic well- distributed purple spots after 3 to 5 min- utes of treatment in 25 cc absolute methyl alcohol acidulated by the addition of 1 to 5 drops of concentrated, chemically pure hydrochloric acid. Transmission : By root grafting, except by root-bark patch grafts, which are inef- fective. Budding and grafting with parts of stem fail to transmit this virus. Thermal inactivation : At 48° C in about 40 minutes in roots. Literature : Hutchins, Georgia State Entomol. Bull., 78, 1933; Phytopath., 29, 1939, 12 (Abst.); Hutchins and Rue, ibid., 29, 1939, 12 (Abst.). 4. Nanus holodisci H. {loc. cit., 127). From New Latin Holodiscus, generic name of ocean spray. Common name: Ocean-spray witches'- broom virus. Host: ROS ACE AE— Holodiscus dis- color Max., ocean spray. Geographical distribution: United States (Oregon and Washington). Induced disease : In ocean-spray, dis- eased branches form clusters of thin wiry shoots with abnormally short internodes and crowded small leaves. Laterals num- erous and more than normally branched. Bronzy red color acquired early by af- fected foliage. Transmission: By aphid. Aphis spir- aeae Schout. (APHIDIDAE). By grafting. Not demonstrated by inocula- tion of expressed juice. Literature: Zeller, Phytopath., 21, 1931, 923-925. 5. Nanus fragariae H. (Holmes, loc. cit., 128; Blastogenus fragariae McKin- ney, Jour. Washington Acad. Science, 34, 1944, 151.) From New Latin Fragaria, generic name of strawberry. Common name: Strawberry witches '- broom virus. Host : ROS ACE AE — Fragaria chiloen- sis Duch. var. ananassa Bailey, culti- vated strawberry. Geographical distribution : United States (western Oregon). Induced disease : In strawberry, leaves numerous, light in color, with spindly petioles, margins of leaflets bent down, runners shortened, plants dwarfed ; flower stalks spindly and unfruitful; root sys- tems normal and well developed. Transmission : By aphis, Myzus fragae- folii Ckll. (APHIDIDAE). Not dem- onstrated by inoculation of expressed juice. Literature : Zeller, Phytopath., 17, 1927, 329-335. 6. Nanus cupuliformans Zeller and Weaver. (Phytopath., 31, 1941, 851.) From diminutive of Latin cupa, tub, and participal from Latin formare, to form. 1208 MANUAL OF DETERMINATIVE BACTERIOLOGY Common name : Strawberry -stunt virus. Host : ROSACEAE — Fragaria chiloen- sis Duch. var. ananassa Bailey, culti- vated strawberry. Geographical distribution : United States (Oregon, Washington). Induced disease : In strawberry, little if any reduction in chlorophyll, plants erect but short ; leaves at first folded, later open, dull in lustre, with papery rattle when brushed by hand, leaflets cupped or with margins turned down, midveins tortuous; petioles I to § normal length; fruits small, usually hard and seedy; roots normal in appearance. Transmission : Bj^ strawberry -leaf aphid, Capitophorus fragaefolii {APHI- DIDAE). By grafting. Not by inocu- lation of expressed juice. 7. Nanus pruni H. {loc. cit., 128). From New Latin Primus, generic name of prune, from Latin prunus, plum tree. Common name: Prune -dwarf virus. Hosts: ROSACEAE— Prnnvs domes- tica L., prune and plum; var. insititia Bailey, the Damson plum, remains symp- tomless. Experimentally, also Prunus persica (L.) Batsch, peach. Insusceptible species: ROSACEAE — Prunus avium L., cherry. Geographical distribution : United States (New York); Canada (British Columbia, Ontario). Induced disease: In prune, leaves small, narrow, rugose, distorted, glazed. Internodes short. Some branches escape and appear normal. Blossoms numerous, mature fruits few. Pistils aborted, pet- als narrow and distorted. In Damson and Bradshaw plums, no obvious mani- festations of disease as a result of infec- tion. Transmission : By budding and other forms of grafting. Not demonstrated by inoculation of expressed juice. No insect vector is known. Literature: Berkeley, Canada, Domin. Dept. Agr., Div. of Bot. and Plant Path., Science Service, Publ. 679, 1941 ; Hilde- brand, Phytopath., 32, 1942, 741-751; Thomas and Hildebrand, Phytopath., 26, 1936, 1145-1148. 8. Nanus sacchari H. {loc. cit., 129). From New Latin Saccharum, generic name of sugar cane, from Latin saccharum, sugar. Common name : Sugar-cane sereh- disease virus. Host : GRAMI NEAE—Saccharum offi- cinarum L., sugar cane. Geographical distribution : Java, Borneo, Sumatra, Moluccas, India, Mau- ritius, Australia, Fiji, Formosa, Hawaii, Ceylon. Induced disease : In sugar cane (Cheri- bon variety), plant dwarfed, shoots stunted, vascular bundles colored by the presence of a red gum ; adventitious roots from many or all nodes. Transmission : Not by inoculation of expressed juice. No insect vector is known . Thermal inactivation : In cuttings of sugar cane, at 52° C in 30 minutes to 1 hour. Infected cane cuttings survive the heat treatment required for cure through inactivation of the causative virus. Literature : Houtman, Arch. Suikerind. Nederland. -Indie, 33, 1925, 631-642; Lyon, Bull. Exp. Sta. Hawaiian Sugar Planters' Assoc, Bot. Ser., 3, 1921, 1-43; Wilbrink, Arch. Suikerind. Nederland. - Indie, 31, 1923, 1-15. Genus V. Rimocortius Milbrath and Zeller. (Phytopath., 32, 1942, 430.) Viruses of the Rough-Bark Group, inducing diseases principally affecting bark, less often wood, leaves, or fruit. Generic name from Latin rima, cleft or fissure, and cortex, bark. The type species is Rimocortius kwanzani Milbrath and Zeller. FAMILY MARMORACEAE 1209 (Note: The genus Citrivir (first named species, Cilrivir psorosis Fawcett, Phyto- path., SI, 1941, 357) was proposed by its author as a genus pro tempore with the avowed purpose of accommodating viruses causing diseases in species of the plant-host genus Citrus. It appears to have been implied by the term genus pro tempore that evidences of natural relationship, when discovered, would permit even the first-named species of this genus to be assigned elsewhere. On the assumption that a permanent genus is nothing more than a type species and such other species as may be added to it by one or another author, it must be felt that a genus pro tempore, however convenient as an expedient, cannot become a permanent genus under any circumstances, because its first -named species would appear not to be a permanent part of the genus and so intended not to be a true type-species. Without a type species there would seem to be no permanent genus concept . The system by which the term Citrivir was coined (explained by its author as use of the genitive of the host-genus name, Citris, plus vir, signifying virus) seems in itself acceptable, for it is commonly agreed that a generic name may be made in an arbitrary manner. It may be noted that use of the stem of the host -genus name (Citr-) with connecting vowel i and suffix -vir, possibly a more orthodox procedure, would have given the same result in the present instance. The original definition of the term Citrivir might be thought to be repugnant as disregarding concepts of natural inter- specific relationships that are essential to the spirit of binomial nomenclature. Were the genus to be regarded as permanent rather than pro tempore, however, the scope of the genus would come to be wholly changed by usage, when, with passage of time related species would be added to what in this case would be a type species, without regard to the unorthodox intent of the original definition but .solely in accordance with similarities between viruses. A generic concept need never be accepted as rigidly defined, whether initially, as has been attempted in this case, or upon further experi- ence, because a genus may still grow by the addition of closely allied new species beyond any limit that may be set. On this account an original, or any subsequent, definition may be regarded as subject to unlimited change so long as the type species is logically retained. The form and definition of the term Citrivir would not, there- fore, militate against its continued use. Its avowedly temporarj'' status alone seems decisively to do so. The originally monotypic genus Rimocortins , publishetl in the following year, was defined only by the combined generic and specific description, and was not referred to a family by its authors. The type, because at first the only species, Rimocortius kwan- zani, is the flowering-cherry rough-bark virus. This type species might well be asso- ciated with the species Citrivir psorosis, citrus-psorosis virus, discussed above, both affecting bark principally, though foliage also to some extent. Although the genus Citrivir was named in 1941 and Rimocortius not until 1942, the first was intended as a temporary assemblage only, as above indicated. It would seem appropriate, there- fore, to include the virus that was known temporarily as Citrivir psorosis in the per- manent genus Rimocortius Milbrath and Zeller and to assign this genus to the family Marinoraceae.) Key to the species of the genus Rimocortius. I. Affecting cherry. 1. Ri7nocortius kwanzani. II. Affecting Citrus. 2. Rimocortius psorosis. III. Affecting pear. 3. Rimocortius pyri. 1210 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Rimocortius kwanzani Milbrath and Zeller. (Phytopath., 32, 1942, 430.) From Kwanzan, name of a variety of flowering cherry. Common name : Flowering-cherry rough -bark virus. Hosts: Prunus serriilata Lindl. var. Kwanzan, flowering cherry; P. avium L., Mazzard cherry. Geographical distribution : United States (Oregon). Induced disease : In flowering cherry, tree dwarfed, deficient in lateral branches; bark deep brown, roughened, splitting longitudinally ; internodes short- ened, bunching leaves; leaves arched downward; midribs of leaves split and cracked on under surface. In Mazzard cherry, no manifestation of disease, but carrier condition ; budded Mazzard stock may transmit disease to healthy Kwanzan cherry cions. Transmission: By budding, generally even if the inserted bud fails to survive. Literature : Milbrath and Zeller, Phy- topath., 32, 1942, 428-430; Thomas, ihid., 32, 1942, 435-436. 2. Rimocortius psorosis (Fawcett) comh. nov. (Citrivir psorosis Fawcett, Phytopath., 31, 1941, 357.) Specific name meaning "of the disease known as psorosis." Common name: Citrus-psorosis virus. Hosts: RUT ACE AE — Citrus sinensis Osbeck, orange; C. limonia Osbeck, lemon; C. maxima Merr., grapefruit. Geographical distribution : World-wide where citrus trees are grown. Induced disease: In citrus, small, elongated, light colored areas or flecks in the region of small veins on young, tender foliage; leaves sometimes warped; (chlo- rotic?) clearing of veins, and chlorotic line patterns, sometimes concentric. Outer layers of bark scale away ; depres- sions and deformities appear in bark and wood. Lemons, as a rule, are more tol- erant than oranges and are not subject to the bark changes. Transmission : By grafting, including root grafting and patch bark grafting. Not by inoculation of expressed juice. No insect vector is known. Literature : Bitancourt et al., Phyto- path., 33, 1943, 865-883; Fawcett, ibid., 21t, 1934, 659-668; Science, 92, 1940, 559- 561; Phytopath., 31, 1941, 356-357; Faw- cett and Bitancourt, ihid., 33, 1943, 837- 864; Rhoads, ibid., 32, 1942, 410-413; Webber and Fawcett, Hilgardia, 9, 1935, 71-109. Strains : Three strains differing from the type have been recognized. The type, var. vulgare Fawcett, Phytopath., 31, 1941, 357, causes psorosis A, the com- mon scaly-bark type of disease, with pustular eruptions of outer layers of bark in limited areas, with or without exuda- tion of gum ; later a drab-gray, cinnamon- drab to rufus discoloration of the wood, accompanied by decline of the affected tree. Others, that contrast with the type, are : 2a. Rimocortius psorosis var. anulatum Fawcett. (Phytopath., 31, 1941, 357.) From Latin anulatus, with a ring. Caus- ing psorosis B, known from California, resembling zonate chlorosis of Brazil in effects on leaves and fruits. Psorosis B is characterized by rapid scaling of outer bark in continuous areas, progressing rapidly along one side of trunk or branch ; gum exudes in advance of scaling, necrosis follows; large circular discolored and corky spots, sometimes concentric, on fruits and mature leaves; on some fruits, circular or semi-circular furrows and bumps ; rapid decline of the affected tree. 2b. Rimocortius psorosis var. concavum Fawcett and Bitancourt. (Phytopath., 33, 1943, 850.) From Latin concavus, concave. Causing concave-gum psorosis, characterized by concavities of various sizes on trunks and larger limbs of af- fected trees, often by zonate patterns on young leaves during periods of rapid growth. FAMILY MARMORACEAE 1211 2c. Rimocortius psorosis var. alveatum Fawcett and Bitancourt. (Phytopath., 33, 1943, 854.) From Latin alveatus, hol- lowed out like a trough. Causing blind- pocket psorosis, characterized by trough- like pockets in bark and wood or bj^ erup- tive lesions. 3. Rimocortius pyri (Holmes) cotnb. nov. {Marmor pyri Holmes, Handb. Phytopath. Viruses, 1939, 76.) From New Latin Pyrus, generic name of pear, from Latin pirus, pear tree. Common name : Pear stony -pit virus. Host : ROSACE AE — Pyrus communis L., pear. Geographical distribution: United States (Oregon, Washington, Califor- nia). Induced disease : In pear, fruit deeply pitted and deformed; bark cracked and resembling oak bark; veinlet chlorosis of some leaves, failure of lateral buds to grow, reduction of foliage. Bartlett and Comice varieties of pear appear to be tolerant, producing sound fruit from infected trees. Transmission: By budding. Xot by inoculation of expressed juice. No insect vector is known. Literature: Kienholz, Phytopath., 29, 1939, 260-267; 30, 1940, 787 (Abst.). Genus VI. Adelonosus Brierley and Smith. (Phytopath., 3^, 1944, 551.) Viruses capable of multiplying in living plants but producing no recognizable symp* toms in these except on interaction with distinct viruses with which they form com- plexes. Transmitted by aphids, by sap, or by both means. Generic name from Greek adelos, invisible, and nosos, disease. Only one species is recognized thus far; this is the type species. Adelonosus lilii Brierley and Smith. 1. Adelonosus lilii Brierley and Smith. (Phytopath., 34, 1944, 551.) From Latin lilium, lily. Common name : Lily-symptomless virus. Host : LILI ACE AE— Lilium longi- florum Thunb., Easter lily. Insusceptible species: All other tested lilies and many related plants in the same and other families (for list, see Phyto- path., 34, 1944,549). Geographical distribution : United States, Japan; probably coextensive with commercial culture of Easter lily. Induced disease : In Easter lily, no obvious manifestation of disease when this virus is present alone ; when together with cucumber-mosaic virus, however, the lily-sj^mptomless virus is a deter- mining factor in the production of ne- crotic-fleck disease; the lily-symptomless virus is so widely distributed in sup- posedly healthy stocks of the Easter lily that cucumber-mosaic virus formerly was thought to be the sole determining factor in necrotic flecking, now recognized to be caused by the virus complex lily- symptomless virus (Adelonosus lilii) plus cucumber-mosaic virus (Marmor cucii- meris); the complex acts independently of the presence or absence of lily latent - mosaic virus (Marmor inite), which is often present with the essential members of the complex in flecked Easter lilies. Transmission : By inoculation of ex- pressed juice, with some difficulty. By aphid. Aphis gossypii Glov., cotton aphid (APHIDIDAE); preinfective period after obtaining virus, 4 to 6 days. 1212 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY III. ANNULACEAE HOLMES. (Handb. Phytopath. Viruses, 1939, 97.) Viruses of the Ringspot Group, causing diseases usually characterized by necrotic; or chlorotic spotting with concentric-ring lesions and eventual recovery from obvious disease with non-sterile immunity. Hosts, higher plants; vectors unknown. There is a single genus. Genus I. Annulus Holmes. {hoc. cit., 97.) Characters those of the family. Generic name from Latin annulus, a ring. The type species is Annulus tahaci Holmes. Key to the species of genus Annulus. I. Found occurring naturally in the Western Hemisphere. A. In tobacco. 1. Annulus tahaci. 2. Annulus zonatus. 3. Annulus orae. 4. Annulus apertus. B. In potato. 5. Annulus dubius. C. In delphinium. 6. Anmtlus delphinii. II. Old World species. 7. Annulus bergerac. 1. Annulus tabaci Holmes. (Handb. Phytopath. Viruses, 1939, 98; Marmor anularium McKinney, Jour. Washington Acad. Sci., S4, 1944, 327.) From New Latin Tahacum, early generic name for tobacco. Common names: Tobacco-ringspot vi- rus, green ringspot virus, yellow ringspot virus, ring spot No. 1 virus. Hosts: SOLA N ACE AE—Nicotiana tahacum L., Petunia violacea Lindl., Solanum tuberosum L. CUCURBITA- CEAE — Cucumis sativus L. Experimen- tally this virus has been found capable of infecting many species of plants in a large number of families ; these include all tested species of the SOLAN ACE AE, SCROPHULARIACEAE, COMPOSI^ TAE, and CUCURBIT ACEAE. Many species of the LEGUMINOSAE are susceptible and one, Vigna sinensis (L.) Endl., is used as an indicator plant for quantitative measurement because it displays conspicuous reddish -brown necrotic lesions around points of initial infection. Geographical distribution: United States. Induced disease : In tobacco, necrotic ring-like primary lesions, followed by secondary necrotic rings on younger leaves. Subsequently, affected plants recover. After recovery from obvious disease, virus content of plants is only 10 to 20 per cent of that of recently infected jjlants. Some varieties may show mosaic-like patterns in young leaves at 16°C. Transmission : By inoculation of e.x- dressed juices. Through about 20 per cent of seeds from diseased petunia plants. Not by dodder, Cuscuta campes- tris Yuncker (CONVOLVULACEAE). Serological relationships : Induces the formation of specific precipitating anti- bodies when injectetl into bloodstream of rabbit. Immunological relationships : Recov- ered tobacco plants are not susceptible to reinfection with this virus but are readily infected with Annulus zonatus or A. orae. This virus produces primary lesions on FAMILY ANNULACEAE 1213 leaves of plants immune to reinfection with A. bergerac. Thermal inactivation : At 68° C in 10 minutes. Filterability : Passes V, X, and perhaps W Berkefeld filters. Other properties: Particle size esti- mated by filtration experiments as about 19 millimicrons. Sedimentation con- stant, 820° = 115 X 10-13 cm. sec.-i dyne~i. Infective in dilutions of 10"^ after purification. Inactivated in 1 hour below pH 3 or above pH 10.8. Recovered plants of tobacco contain 0.002 mg of virus per gram, recently infected plants about 6 times as much. Optimum con- ditions for retaining infectivity of stored virus include suspension in 0.01 M phos- phate buffer at pH 7 and storage at 4° C. Literature: Fenne, Phytopath., 21, 1931, 891-899; Fromme et al., ibid., 17, 1927, 321-328; Henderson, ibid., 21, 1931, 225-229; Henderson and Wingard, Jour. Agr. Res., J,3, 1931, 191-207; Price, Con- trib. Boyce Thompson Inst., I^, 1932, 359-403; Phytopath., 26, 1936, 503-529; Am. Jour. Bot., 27, 1940, 530-541; Am. Naturalist, 74, 1940, 117-128; Priode, Am. Jour. Bot., 15, 1928, 88-93 ; Stanley, Jour. Biol. Chem., 129, 1939, 405-428, 429-436; Stanley and Wyckoff, Science, 85, 1937, 181-183; Valleau, Kentucky Agr. Exp. Sta., Bull. 327, 1932; Wingard, Jour. Agr. Res., 37, 1928, 127-153; Woods, Contrib. Boyce Thompson Inst., 5, 1933, 419-434. Strains: A number of distinctive strains have been collected in nature and studied experimentall}'. The following have been given varietal names to dis- tinguish them from the type, var. vir- giniensis H., loc. cit., 98: la. Annulus tabaci var. kentuckiens is H. {loc. cit., 99). Differing from the typical strain in producing less necrosis and less stunting in tobacco. (Price, Phytopath., 26, 1936, 665-675; Valleau, Kentucky Agr. Exp. Sta., Bull. 327, 1932.) lb. Annulus tabaci var. avratus H. {loc. cit., 100). Secondary lesions in tobacco at first yellow spots or rings, be- coming necrotic subsequently. Recov- ery less complete than with type, abnor- mal yellowing of old leaves tending to persist. (Chester, Phytopath., 25, 1935, 686-701; Price, Phytopath., 26, 1936, 665-675; Valleau, Kentucky Agr. Exp. Sta., Bull. 327, 1932; Phytopath., 29, 1939, 549-551.) 2. Annulus zonatus H. {loc. cit., 101 j. From Latin zonatus, zonate. Common names : Tomato -ringspot vi- rus, ring spot No. 2 virus. Hosts : SOLAN AC EAE—Nicotiana tabacum L., tobacco. Experimentally this virus has been found to infect many species of plants in a large number of families. Geographical distribution : United States. Induced disease : In tobacco, zonate necrotic primary lesions and, temporarily, secondary lesions of the same type; recovery with specific, non-sterile im- munity. In tomato, systemic infection, yellowish-green or necrotic ring-like le- sions; stunting. Transmission: By inoculation of ex- ])ressed juice. Immunological relationships : Recov- ered plants are immune to reinfection but are still susceptible to Annulus tabaci, A. bergerac, and several mosaic -type viruses that have been tested. Thermal inactivation : At 55 to 60° C in 10 minutes. Filterability : Passes Gradocol mem- brane 100 millimicrons in average pore diameter. Particle size estimated as 50 millimicrons or less. Literature : Price, Phytopath., 26, 1936 665-675 ; Am. Jour. Bot., 27, 1940, 530-541 . 3. Annulus orae H. {Holmes, loc. cit., 103; Tractns orae Valleau, Phytopath., 30, 1940, 826.) From Latin ora, edge, in reference to occurrence of induced dis- ease near edge of tobacco fields. Common name : Tobacco-streak virus. Hosts : SOLA NA CEAE— Nicotiana tabacum L., tobacco. Experimentally, a 1214 MANUAL OF DETERMINATIVE BACTERIOLOGY number of other solanaceous plants have been reported as susceptible, but not Capsicvm fruiescens L., pepper; Lyco- persicon esculentvm Mill., tomato; So- larium melongena L., eggplant; or S. tuberosum L., potato. Geographical distribution : United States. Induced disease : In tobacco, local and systemic necrosis in 3 daj's, with irregular spot, line, and ring-like lesions, followed by recovery from necrotic mani- festations of disease. Recovered leaves may show a mild mottling and regularly contain virus ; reinoculation does not induce formation of necrotic lesions in them. Transmission : By inoculation of ex- pressed juice. Not through seeds from diseased plants. Immunological relationships : No cross- protection with respect to A. tabaci, and several viruses of the mosaic group. Thermal inactivation : At 53° C in 10 minutes. Literature : Johnson, Phytopath., 26, 1936, 285-292; Trans. Wisconsin Acad. Sciences, Arts and Letters, SO, 1937, 27-34. 4. Annulus apertus spec. nov. From Latin apertus, frank. Common name: Broad-ringspot virus. Hosts : SOLAN ACE AE—Nicotiana tabacum L., tobacco. Experimentally also to many species in this and other families. Insusceptible species: CHENOPODI- ACEAE—Beta vulgaris L. C UC URBI- TACEAE — Ciirullus vulgaris Schrad. LEGUMINOSAE—Medicago saliva L., Melilotus alba Desr. Geographical distribution : United States (Wisconsin). Induced disease : In tobacco, indistinct yellow-spot primary lesions, becoming chlorotic or necrotic rings with concen- tric markings; small chlorotic rings, sometimes concentric, or fine brown ne- crotic rings as secondary lesions ; young leaves puckered at first, somewhat mal- formed. Transmission : By inoculation of ex- pressed juice. Immunological relationships : Protects against reinfection with homologous virus but leaves host susceptible to infection by Annulus tabaci, A. zonatus, and some mosaic -type viruses. Literature : Johnson and Fulton, Phy- topath., 32, 1942, 605-612. 5. Annulus dubius (Holmes) comb.nov. (Marmor dubium H., loc. cit., 42.) From Latin dubius, uncertain, in reference to a common name, potato virus X, often used to designate this virus. Common name : Potato-mottle virus (strains of this virus have been studied at various times under the names potato latent virus, potato virus X, potato- anecrosis virus, virulent latent virus, simple mosaic virus, healthy potato virus, Hyoscyamus IV virus, President streak virus, potato foliar-necrosis virus, potato acronecrotic streak virus, Up-to-Date streak virus, potato viruses B and D, Solanum viruses 1, 4, and 6.) Hosts: SOLA N ACE A E— Solanum tu- berosuni L., potato; Lycopersicon esculen- tum Mill., tomato. Experimentally, also SOLAN ACE AE — Capsicum frutescens L., pepper; Datura stramonium L., Jim- son weed ; Hyoscyamus niger L., henbane ; Nicotiana tabacum L., tobacco; Physalis alkckengi L. ; Solanum didcamara L., bit- tersweet ; S. nigrum L., black nightshade. AM ARAN TH ACE A E— Amaranthxis retroflexus L. COM POSIT AE— Chrysan- themum morifolium Ram. SCROPHU- LARIACEAE — Veronica sp., common speedwell. Geographical distribution : Widespread throughout the world; present in all known stocks of tubers of some potato varieties in the United States. Induced disease : In potato, usually no chlorotic mottling, sometimes a little ; intracellular inclusions of the vacuolated, granular type ; some varieties that are virtually immune in the field owe their tendency to localize the virus in necrotic primary lesions or in top-necrosis of first systemically infected plants to a dom- FAMILY ANNULACEAE 1215 inant allele of a gene nx, which charac- terizes plants showing a mosaic of some degree of intensity on infection with this virus ; the variety known as S41956 is im- mune to all tested strains of the virus and possesses two dominant genes both re- quired for resistance. In tomato, sys- temic mild chlorotic mottling ; if a strain of tobacco-mosaic virus is also present, a severe systemic necrosis, known as double-virus streak, is induced. Transmission : By inoculation of ex- pressed juice. Experimentally, by leaf contacts mainly under the influence of wind. No insect vector is known. Not transmitted through true seeds of the potato. Serological relationships : Cross precip- itin reactions between constituent strains of this virus. No cross reaction with potato aucuba-mosaic, potato mild-mo- saic, potato -veinbanding, tobacco-mo- saic, tobacco-etch, tobacco-ringspot or pea-mosaic virus. Antisera prepared by injecting rabbits intravenously with virus inactivated by nitrous acid, like those prepared with active virus, fix comple- ment and flocculate with virus suspen- sions (though not with juice of healthy host plants); they are also effective in neutralizing the virus. Immunological relationships : Tobacco and Datura plants infected by the type strain of this virus become immune to the more severe potato-ringspot strain. No protection against the severe strain is af- forded by previous infection with to- bacco-mosaic, tobacco-ringspot, tomato spotted-wilt, or cucumber-mosaic virus. Thermal inactivation : At 70° C. in 10 minutes. Filterability : Passes Pasteur-Cham- berland Li, L3, and L5 filters. Other properties : Digested by 0.02 per cent solution of pepsin in 3 hours at pH 4, at 38° C. Digested also by trypsin. Inactivated by papaine and cyanide, but by neither separately. Isoelectric point near pH 4. Dilute solutions show anisot- ropy of flow. Concentrated solutions are spontaneously birefringent. Proper- ties of the type strain have been less studied than those of the potato-ring- spot strain of this virus. Literature: Bawden, Proc. Roy. Soc. London, Ser. B, 116, 1934, 375-395; Baw- den and Pirie, Brit. .Jour. Exp. Path., 17, 1936, 64-74 ; Bawden et al., ibid., 17, 1936, 204-207; Blodgett, Phytopath., 17, 1927, 775-782; Bohme, Phytopath. Ztschr., 6, 1933, 517-524; Cadman, Jour. Genetics, U, 1942, 33-52; Chester, Phytopath., 27, 1937, 903-912; Clinch, Sci. Proc. Roy. Dublin Soc, 23, 1942, 18-.34; Johnson, Wisconsin Agr. Exp. Sta., Res. Bull. 63, 1925; Koch, Phytopath., 23, 19.33, 319- 342; Kohler, Phytopath. Ztschr., 5, 1933, 567-591; 7, 1934, 1-30; Loughnane and Murphy, Nature, I4I, 1938, 120; van der Meer, Cent. f. Bakt., II Abt., 87, 1932, 240-262; Salaman, Nature, 131, 1933, 468; Schultz et al., Phytopath., 27, 1937, 190- 197; 30, 1940, 944-351 ; Spooner and Baw- den, Brit. .Jour. Exp. Path., 16, 1935, 218-230; Stevenson et al., Phytopath., 29, 1939, 362-365. Strains : Several variants of potato- mottle virus, differing from the type, var. vulgaris H. {loc. cit., 42), have been recog- nized as distinctive varieties under the following names : 5a. Annulus dubius var. annulus H. {loc. cit., 44). From Latin annulus, ring. Common name : Ringspot strain of potato-mottle virus. Necrotic primary and secondary ring-like lesions in experi- mentally infected tobacco plants. Indis- tinguishable from type strain by ordinary precipitin test, but distinguishable when appropriately absorbed sera are used. This strain has been more frequently studied than the type. Juice of infected tobacco plants contains about 0.02 to 0.10 mg of virus per ml. Sedimentation constants, 820° = 113 X lO-i^ and 131 X 10~i^ cm. sec.~i dyne"^ -Dissymmetry constant 2.78. Molecular weight 26 X 10". Particle size estimated to be 433 by 9.8 millimicrons, 43.9 times as long as wide. Isoelectric point near pH 4. Stable between pH 4 and pH 9.5. Con- centrated solutions are spontaneously birefringent. Dilute solutions show 121G MANUAL OF DETERMINATIVE BACTERIOLOGY anisotropyof flow. Destroyed by drying. Inactivated bj' papaine and cyanide, but by neither separately. Digested by 0.02 per cent solution of pepsin in 3 hours at pH 4, at 38° C. Digested also by trypsin. About 6 per cent of the purified virus is reported to be a pentose nucleic acid, but the carbohydrate to phosphorus ratio is about twice that for yeast nucleic acid. Guanine and pentose present. Analysis of sedimented virus, carbon 47.7 to 49.5 per cent, hydrogen 6.8 to 7.7 per cent, nitrogen 14.6 to 17.0 per cent, phosphorus 0.4 to 0.7 per cent, sulfur 1.1 per cent, carbohydrate 2.5 to 4.3 per cent, ash 2.0 to 2.5 per cent. Reduction of carbo- hydrate content of sample to 2.5 per cent does not reduce activity of virus ; further reduction inactivates. (Ainsworth, Ann. Appl. Biol., 21, 1934, 581-587; Bawden, Brit. Jour. Exp. Path., 16, 1935, 435-443; Bawden and Pirie, ibid., 19, 1938, 66-82; Birkeland, Bot. Gaz., 95, 1934, 419-436; Chester, Phytopath., S6, 1936, 778-785; Johnson, Wisconsin Agr. Exp. Sta., Res. Bull. 76, 1927; Loring, Jour. Biol. Chem., 126, 1938, 455-478; Loring and Wyckoff, ibid., 121, 1937, 225-230.) 5b. Annulns diibius vay . flavus Yl. {loc. cit., 46). From Latin flavus, yellow. Common name : Yellow-mottle strain of potato-mottle virus. Differing from the type by imparting a yellow cast to foliage of infected potatoes. (Putnam, Canad. Jour. Res., Sec. C, 15, 1937, 87- 107.) 5c. Annulus dubiiis var. obsciirus H. (loc. cit., 46). From Latin obscurus, ob- scure. Common name : Masked-mottle strain of potato-mottle virus. Differing from the type by systemically infecting potato, tobacco, and Jimson weed with- out symptoms under ordinary experi- mental conditions; in pepper, however, systemic necrosis is induced, as by all known strains. (Chester, Phytopath., 26, 1936, 778-785.) 6. Annulus delphinii spec. nov. From New Latin Delphinium, generic name of host . Common names : Delphinium-ringspot virus, perennial -delphinium ringspot virus. Hosts: RAN UNCULACEAE— Del- phinium sp., perennial delphiniums. Experimentally, also to CHENOPODI- ACEAE—Beta vulgaris L. CUCUR- BIT ACEAE — Cucumis sativus L., cu- cumber. MALVACEAE — Gossypium hirsutum L. RANUNC UL ACEAE— Ranunculus asiaticus L. (symptomless carrier). SOL AN ACEAE— Datura stramonium L., Nicotiana alata Link and Otto, N . gluiinosa L., N. rustica L., N. tabacum L., Petunia hybrida Vilm. Geographical distribution: United States (California). Induced disease : In perennial del- phiniums, faint chlorotic rings with green or yellow centers on young leaves ; irregular chlorotic spots, yellow bands, or irregular chlorotic rings on mature leaves. Transmission : By inoculation of ex- pressed juice in the presence of finely powdered carborundum. Thermal inactivation : At 65° C in 10 minutes. Literature : Severin and Dickson, Hil- gardia, 14, 1942, 465-490. 7. Annulus bergerac H. (loc. cit., 102). From Bergerac, a town in southwest France. Common name : Bergerac-ringspot virus. Hosts : SOLAN ACEAE— Nicotiana tabacum L., tobacco. Experimentally, this virus has been transferred to several other solanaceous plants and to Phaseolus vulgaris L., bean, in the family LEGUM- INOSAE. Geographical distribution: France. Induced disease : In tobacco, thin necrotic -ring primary lesions, followed by FAMILY ANNULACEAE 1217 systemic mottling with some chlorotic rings on the dark green islands. Later, complete recovery occurs, with non- sterile immunity. Transmission : By inoculation of ex- pressed juice. Immunological relationships : Recov- ered plants are susceptible to infection by Annulus tab aci and A. zonatus. Thermal inactivation : At 80° C in 10 minutes. Literature : Smith, A textbook of plant virus diseases, P. Blakiston's Son and Co., Inc., Philadelphia, 1937, 285-289. 1218 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY IV. RUGACEAE HOLMES. (Handb. Phytopath. Viruses, 1939, 114) Viruses of the Leaf -Curl Group, causing diseases characterized by suddenly arrested development of invaded tissues, resulting in leaf curl, enations, and other deformities. Vectors, typically white-flies (ALEYRODIDAE). There is a single genus. Gentis I. Ruga Holmes. {Loc. cit., 114.) Characters those of the family. Generic name from Latin ruga, a wrinkle. The type species is Ruga tabaci Holmes. Keij to the species of genus Ruga. I. Infecting tobacco. II. Infecting cotton. III. Infecting cassava (Manihot). IV. Infecting sugar-beet. 1. Ruga tabaci Holmes. (Handb. Phy- topath. Viruses, 1939, 115.) From New Latin Tabacum, former generic name of tobacco. Common names : Tobacco leaf -curl virus, kroepoek virus, curl-disease virus, crinkle-disease virus. Hosts: SOLAN ACE AE—Nicotiana tabacum L., tobacco. COMPOSITAE— Vernonia iodocalyx, V. cineria, Ageratum conyzoides L., Synedrella nodi flora Gaertn . Experimentally, also other solanaceous plants. Insusceptible species: MALVACEAE — Gossypium hirsutum L., cotton. Geographical distribution : Tanganyika, Southern Rhodesia, Southern Nigeria, Nyasaland, India, Sumatra, Formosa. Induced disease : In tobacco, leaves curled and crinkled, with occasional leafy outgrowths or enations. Veins greened and thickened. No chlorosis nor necro- sis. Plant stunted. Transmission: By white-fly, Bemisia gossypiperda Misra and Lamba (ALEY- RODIDAE). By grafting. Not by in- oculation of expressed juice. Literature : Kerling, Phytopath., £3, 1933, 175-190; Mathur, Indian Jour. Agr. 1. Ruga tabaci. 2. Ruga gossypii. 3. Ruga bemisiae. I 4. Ruga verriicosans . Sci., 3, 1933, 89-96; Matsumoto and Tateoko, Trans. Nat. Hist. Soc. Formosa, 30, 1940, 31-33; Pal and Tandon, Indian Jour. Agr. Sci., 7, 1937, 363-393; Pruthi and Samuel, ibid., 7, 1937, 659-670; Storey, Nature, 128, 1931, 187-188; East African Agr. Jour., 1, 1935, 148-153; Thung, Meded. Proefsta. Vorstenl. Ta- bak Java, 72, 1932; 78, 1934. 2. Ruga gossypii H. (loc. cit., 116). From Latin gossypium, cotton. Common names : Cotton leaf -curl virus, cotton leaf -crinkle virus. Hosts: MALVACEAE— Gossypium hirsutum L., cotton ; G. peruvianum Cav. ; G. vitifolium Lam.; Hibiscus cannabinus L. ; H. esculentus L. ; //. sabdariffa L. ; Althaea rosea Cav., hollyhock; Sakel (hybrid) cotton. Geographical distribution : The Sudan and Nigeria, in Africa. Induced disease : In cotton, clearing of veins, blistering and pale spotting of leaves ; leaves puckered at edge and un- symmetrical. Internodes shortened, producing bunchy growth. Transmission : By white-fly, Bemisia gossypiperda Misra and Lamba (ALEY- FAMILY RUGACEAE 1219 RODIDAE). Not through egg of insect vector. Not by inoculation of expressed juice. Not through soil. Not through seeds from diseased plants. Literature : Bailey, Empire Cotton Growing Rev., 11, 193-4, 280; Kirkpatrick, Bull. Entom. Res., 21, 1930, 127-137; 22, 1931, 323-363. 3. Puga bemisiae H. (Holmes, loc. cit,., 117; Ochrosticta bemisiae McKinney, Jour. Washington Acad. Sci., 34, 1944, 149.) From New Latin Bemisia, generic name of insect vector. Common names : Cassava-mosaic virus, cassava Krauselkrankheit virus. Hosts: EUPHORBIACEAE—Mani- hot utilissima Pohl, cassava; M. palmata Muell.; M. dulcis. Geographical distribution: Gold Coast, Belgian Congo, French Cameroons, Rho- desia, Liberia, Madagascar, probably throughout Africa and adjacent islands; Java. Induced disease : In Manihot utilissima, leaves unsymmetrical, curled, distorted, mottled; internodes shortened, plants stunted. Axillary buds produce an extra number of side branches. Transmission: By white-flies (ALEY- RODIDAE), Bemisia nigeriensis Corb., in Southern Nigeria, and B. gossypiperda Misra and Lamba, in Belgian Congo and Tanganyika. White-flies infect only young leaves. Not by needle-puncture, rubbing, or hypodermic-needle injection of juice expressed from diseased plants. Literature : Dade, Yearbk. Dept. Agr. Gold Coast, 1930, 245; Dufrenoy and Hedin, Rev. Bot. Appl., 9, 1929, 361-365; Golding, Trop. Agric, Trinidad, 13, 1936, 182-186; Kufferath and Ghesquiere, Compt. rend. Soc. Biol. Beige, 109, 1932, 1146; Lefevre, Bull. Agr. Congo Beige, 26, 1935, 442; McKinney, Jour. Agr. Res., 39, 1929, 557-578; Muller, Bull. Inst. Plantenziekt., 24, 1931, 1-17; Pascalet, Agron. Colon., 21, 1932, 117; Staner, Bull. Agr. Congo Beige, 22, 1931, 75; Storey, East Afr. Jour., 2, 1936, 34-39 ; Storey and Nichols, Ann. Appl. Biol., 25, 1938, 790- 806; Zimmermann, Pflanzer, 2, 1906, 145. 4. Ruga verrucosans Carsner and Ben- nett. {Chlorogenus eutetlicola (in error for eutettigicola, from New Latin Eutettix, genus name of a vector, and Latin -cola, dweller in or inhabitant of) Holmes, 1939, loc. cit., 11 ; Carsner and Bennett, Science 98, 1943, 386.) From Latin, meaning: causing rough swellings. Common name : Sugar-beet curly-top virus. Hosts : Very wide range in many fami- lies of dicotyledonous plants. Among the horticulturally important host plants are the sugar beet {Beta vulgaris L., CHENOPODIACEAE); bean {Phase- oliis vulgaris L., LEGUMINOSAE); squash {Cucurbita species, CUCURBI- TACEAE) ; and tomato {Ly coper sicon esculcntumUiW., SOLAN ACE AE). Geographical distribution : Western North America; in Argentina a strain of virus thought to belong here has been reported but has not yet been fully de- scribed. Induced disease : In beet, clearing of veins, leaf curling, sharp protuberances from veins on lower surface of leaves, increase in number of rootlets, phloem degeneration followed by formation of supernumerary sieve tubes, retardation of growth. In tomato, (western yellow blight or tomato yellows), phloem de- generation followed by formation of su- pernumerary sieve tubes, retardation of growth, dropping of flowers and buds, rolling, yellowing and thickening of leaves, root decay, usually followed by death, sometimes by recovery. Occasionally there is relapse after recovery. In cucur- bitaceous plants, stunting, bending up- ward of tip of runner, yellowing of old leaves, abnormally deep green in tip leaves and stem; Marblehead squash is tolerant, showing only mild witches' broom formation and phyllody. In bean, infected when young, thickening and downward curling of first trifoliate leaf, which becomes brittle and will break easily from the stem; leaves become chlorotic, plant stops growing and usually dies soon; older plants are also suscepti- ble to infection; they may survive until 1220 MANUAL OF DETERMINATIVE BACTERIOLOGY the end of the season, showing puckering and downward curling of leaves at the top of the plant, reduction in size of new leaves, and shortened internodes, or they may gradually become chlorotic and die. Transmission : By leafhopper, Eutettix tenellus (Baker) (CICADELLIDAE) with 4 to 12 hour preinfective period. Through dodder, Cuscuta carnpestris Yuncker (CONVOLVULACEAE). Not, with any regularity at least, by mechani- cal inoculation of expressed juice. Not through seeds of diseased plants to seed- lings germinating from them. The leaf- hopper, Agalliana ensigera Oman {CI- CADELLIDAE), is said to transmit a South American strain of sugar-beet curly-top virus, but evidence for identity of the virus has not yet been reported in detail. Thermal inactivation : At 75° to 80° C in 10 minutes. Filterability : Passes Berkefeld V, N, and W, Mandler medium and fine, and Chamberland Li, Lg, Lo, Ly, L9, Ln and Li3 filters. Other properties : Withstands alcohol and acetone treatments. A pH of 2.9 or lower inactivates, but an alkaline reac- tion as high as pH 9.1 does not inactivate, in 2 hours. Virus active after at least 8 years in tissues of thoroughly dried young sugar-beet plants, 6 months in dried leaf- hoppers, and 10 months in dried phloem exudate. Strains : In general it has proved possi- ble to modify strains by host passage, some hosts like Chenopodium murale L. appearing to select less virulent strains, others like Stellaria media (L.) Cyr. reversing this selection and restor- ing virulence. Literature : Bennett, Jour. Agr. Res., 48, 1934, 665-701; 50, 1935, 211-241; 56, 1938, 31-52; Phytopath., 32, 1942, 826- 827; Carsner, Phytopath., 15, 1925,745- 757; U. S. Dept. Agr., Tech. Bull. 360, 1933; Jour. Agr. Res., 33, 1926, 345-348; Dana, Phytopath., 28, 1938, 649-656; Fawcett, Re vista Industrial y Agricola de Tucumdn, 16, 1925, 39-46; Fife, Phyto- path., 30, 1940, 433-437; Giddings, Phy- topath., 27, 1937, 773-779 ; Jour. Agr. Res., 56, 1938, 883-894; Lackey, Jour. Agr. Res., 55, 1937, 453-460; Lesley and Wallace, Phytopath., 28, 1938, 548-553; Murphy, ibid., 30, 1940, 779-784; Severin, Hilgardia, 3, 1929, 595-636; Severin and Freitag, ibid., 8, 1933, 1-48; Severin and Henderson, Hilgardia, 3, 1928, 339-393; Severin and Swezy, Phytopath., 18, 1928, 681-690; Shaw, U. S. Dept. Agr., Bull. 181, 1910. FAMILY SAVOIACEAE 1221 FAMILY V. SAVOIACEAE HOLMES. (Handb. Phytopath. Viruses, 1939, 131.) Viruses of the Savoy-Disease Group, causing diseases characterized mainly by crinkling of foliage. Vectors, true bugs (PIESMIDAE and MIRIDAE). There is a single genus. Genus I. Savoia Holmes. {hoc. cit., 131.) Characters those of the family. Generic name from French chou de Savoie, cabbage of Savoy, a cabbage with wrinkled and curled leaves. The type species is Saroia betae Holmes. Key to the species of genus Savoia. I. Infecting beet. II. Infecting rape and rutabaga. 1. Savoia betae Holmes. (Handb. Phytopath. Viruses, 1939, 132.) From Latin beta, beet. Common names : Beet-Krauselkrank- heit virus, sugar-beet leaf-curl virus, sugar-beet leaf -crinkle virus, Kopfsalat virus. Host : CHE NOPODI ACE AE— Beta vulgaris L., beet. Geographical distribution: Germany, Poland. Induced disease: In beet, veins of leaves swollen, retarded in growth, caus- ing crinkling. New leaves remain small and incurved, forming a compact head. Old leaves die ; plant succumbs before harvest time. Pre patent period in plant , 3 to 9 weeks. Transmission : By tingid bug, Piesma quadrataYieh. (PIESMIDAE). Not by inoculation of expressed juice. Literature : Wille, Arb. Biol. Reich- sanst. Land- u. Forstw.; 16, 1928, 115-167. 2. Savoia piesmae H. {lac. cit., 132). From New Latin Piesma, generic name of insect vector. Common name : Beet -savoy virus. Host: CHENOPODI ACE AE— Beta vulgaris L., beet. 1. Savoia betae. 2. Savoia piesmae.. 3. Savoia napi. Geographical distribution : United States (Michigan, Ohio, Minnesota, Ne- braska, South Dakota, Colorado, Wyom- ing) and Canada. Induced disease : In beet, leaves dwarfed, curled down, small veins thick- ened. Phloem necrosis in roots. Pro- dromal period in plant, 3 to 4 weeks. Transmission : By tingid bug, Piesma cinerea (PIESMIDAE). Not by inocu- lation of expressed juice. Literature: Coons et al., Phytopath., 27, 1937, 125 (Abst.); Hildebrand and Koch, ibid., 32, 1942, 328-331. 3. Savoia napi H. (loc. cit., 133). From New Latin Napus, former generic name of rape, Brassica napus L. Common name : Rape -savoy virus. Hosts: CRUCIFERAE — Brassica na- pus L., rape ; B. napobrassica Mill., ruta- baga. Geographical distribution : Germany. Induced disease : In rape, twisting and crinkling of young leaves ; premature death of old leaves and of plants ; in sur- viving plants, inhibition of growth in spring. In rutabaga, mottling and crink- ling of leaves, with formation of fissures at leaf edges. Plants rarely killed. 1222 MANUAL OF DETERMINATIVE BACTERIOLOGY Transmission : By inoculation of ex- pressed juice. By the tarnished plant bug, Lygus pratensislAnn.{MIRIDAE). The insect vector retains this virus dur- ing intervals between crops. Literature : Kaufmann, Arb. Biol. Reichsanst. Land- u. Forstw., 21, 1936, 605-623; Mitteil. Landwirtsch., 37, 1936; Pape, Deutsch. Landwirtsch. Presse, 26, 1935. FAMILY LETHACEAE 1223 FAMILY VI. LETHACEAE HOLMES. (Handb. Phytopath. Viruses, 1939, 135.) Virus strains of the Spotted-Wilt Group, causing diseases characterized by bronzing of foliage, streaking of stems, blighting of tips, necrotic spotting of foliage higher plants; vectors, thrips (THRIPIDAE). There is a single genus Hosts, Genus I. Lethiun Holmes. (Loc. cit., 135.) Characters those of the family. Generic name from Latin lethum, death. At present there is but one known species, though this is reported to be nearly world-wide in distribution. In some areas it may have been confused with entities needing sep- arate recognition. The t3'pe species is Leihum australiense Holmes. 1. Lethum australiense Holmes {loc. cit., 136). From Australia, where virus was first described. Common names : Tomato spotted-wilt virus, kromnek or Kat River disease virus. Also, pineapple yellow-spot or side-rot virus (according to Sakimura, Phytopath., 30, 1940, 281-299). Hosts : Very numerous species in manj^ families of higher plants. Among those most often noted are : SOLAN ACE AE— Lycopersicon esculentum Mill., tomato; Nicotiana tabacum L., tobacco; Solanutn tuberosum L., potato. COMPOSITAE — Lactuca saliva L., lettuce. LEGUM- INOSAE—Pisum sativum L., pea. BROMELI ACE AE— Ananas comosus Merr., pineapple. Geographical distribution: Australia, British Isles, United States, South Africa Hawaii, Xew Zealand, Europe, China, South America. Induced disease : In tomato, bronze ring-like secondary lesions, plant stunted, some necrosis; later yellowish mosaic with some leaf distortion. Fruit fre- quently marked with concentric rings of pale red, yellow, or white. In tobacco, primary necrotic lesions followed by sys- temic necrosis, with stem streak, crook- neck, often stunting with subsequent wilting and death, sometimes temporary recovery followed by recurrence of sys- temic necrosis. In lettuce, plant yel- lowed, retarded in growth; brown blem- ishes in central leaves, affected spots dying, becoming like parchment but with brown margins. Axillary shoots may show chlorotic mottling. In pea, pur- plish necrotic streaks on stem; at first, leaves mottled; later, necrotic spots damage foliage; pods show circular ne- crotic spots or wavy lines, or, if severely affected, may collapse; seeds may show necrotic lesions. In potato, zonate ne- crotic spots on upper leaves, necrotic streaks on stems; stems collapse at top; plant is stunted, yield of tubers small. In pineapple, at first an initial spot or primary lesion | to * inch in diameter, raised, yellowish, on upper surface of young leaf; later chlorotic spotting of young leaves, crook-neck because of necrotic foci in stems and fruits (side rot) ; plant may die. Transmission : By inoculation of ex- pressed juice; the addition of fine car- borundum powder to inoculum facilitates transmission bj' rubbing methods. By thrips, Frankliniella lycopersici Andre- wart ha (formerly identified as F. insu- laris Franklin), F. occidentalis Perg., F. moultoni Hood, and F. schultzei (Try- bom) {THRIPIDAE). Also by Thrips tabaci Lind. {THRIPIDAE). In F. lycopersici, thrips must pick up virus while still a nymph; virus persists through pupation and emergence as adult ; preinfective period in vector, 5 to 9 days. Virus is not transmitted through eggs of 1224 MANUAL OF DETERMINATIVE BACTERIOLOGY infective thrips. Probabl}' not through seeds of infected plants. Not through soil. Immunological relationships: Infects tobacco plants previously infected with tobacco-mosaic, potato-mottle, tobacco- ringspot, and tomato-ringspot viruses. Thermal inactivation : At 42° C in 10 minutes. Filterability : Passes Gradocol mem- brane of 0.45 micron pore diameter. Other properties : Virus readily inacti- vated by desiccation or by action of oxi- dizing agents ; activity prolonged by pres- ence of sodium sulfite, cystein, or by low temperatures. Unstable at pH values below 6 and above 9. Literature: Ainsworth et al., Ann. Appl. Biol., 21, 1934, 566-580; Andrewar- tha, Trans. Roy. Soc. of So. Australia, 61, 1937, 163-165; Bald and Samuel, ibid., 21, 1934, 179-190; Berkeley, Scientific Agr., 15, 1935, 387-392; Best, Austral. Chem. Inst. Jour, and Proc, 4, 1937, 375-392; Best and Samuel, Ann. Appl. Biol., 23, 1936, 509-537; 759-780; Carter, Phyto- path., 29, 1939, 285-287; Lewcock, Queensland Agr. Jour., 4S, 1937, 665-672; Linford, ibid., 22, 1932, 301-324; Magee, Agr. Gaz. of New South Wales, ^7, 1936, 99-100, 118; McWhorter and Milbrath, Phytopath., 25, 1935, 897-898 (Abst.); Oregon Agr. Exp. Sta., Circ. 128, 1938; Milbrath, Phytopath., 29, 1939, 156-168; Moore, Nature, H7, 1941, 480-481 ; Moore and Anderssen, Union of So. Africa, Dept. Agr., Science Bull. 182, 1939; Parris, Phytopath., 30, 1940, 299-312; Rawlins and Tompkins, ibid., 26, 1936, 578-587; Sakimura, ibid., 30, 1940, 281-299; Sam- uel and Bald, Ann. Appl. Biol., 20, 1933, 70-99; Jour. Agr. So. Australia, 37, 1933, 190-195; Samuel et al., Counc. Scient. Indus. Res., Austral., Bull. 44, 1930; Ann. Appl. Biol., 22, 1935, 508-524; Shapovalov, Phytopath., 24, 1934, 1149 (Abst.); Smith, Nature, 127, 1931, 852- 853; Ann. Appl. Biol., 19, 1932, 305-330; Jour. Minist. Agr., 39, 1933, 1097-1104; Jour. Roy. Hort. Soc, 60, 1935, 304-310; Snyder and Thomas, Hilgardia, 10, 1936, 257-262 ; Takahashi and Rawlins, Phyto- path., 24, 1934, 1111-1115; Taylor and Chamberlain, New Zealand Jour. Agr., 54, 1937, 278-283; Whipple, Phytopath., 26, 1936, 918-920. Strains : A strain differing somewhat from the type, var. typicum H. {loc. cit., 136), has been described as damaging tomatoes in the northwestern United States. It has been given a distinctive varietal name : la. Lelhum aiistraliensc var. lethale H. (loc. cit., 138). From Latin lethalis, deadly. Common names : Tip-blight strain of tomato spotted-wilt virus, Oregon tip-blight virus, tomato die-back streak virus, tomato tip-blight virus. Differs from the type in causing necrotic leaf spotting, stem streaking, and tip blighting in most hosts, without mottling or bronzing of foliage ; yet in Tropaeolurn inajiis L., there is little necrosis. In tomato, systemic necrosis, terminal shoots blighted and blackened; dead tips stand upright above living foliage. Fruits rough and pitted, with internal pockets of necrotic tissue or with sub- epidermal necrosis, appearing externally as concentric brown bands. (McWhorter and Milbrath, Oregon Agr. Exp. Sta., Circ. 128, 1938; Milbrath, Phytopath., 29, 1939, 156-168.) FAMILY BORRELINACEAE 1225 Suborder III. Zoophagineae suhordo novus. From Greek phagein, to eat, and zoon, an animal. Viruses infecting animals but having no plant hosts, so far as known. Key to the families of suborder Zoophaginae. 1. Inducing diseases in insects as exclusive hosts. Family I. Borrelinaceae, p. 1225. 2. Inducing diseases of the pox group. Famil}'' II. Borreliotaceae, p. 1229. 3. Inducing diseases of the encephalitis group. Family III. Erronaceae, p. 1248. 4. Inducing diseases of the yellow-fever group. Family IV. Charonaceae, p. 1265. 5. Inducing diseases of the infectious anemia group. Family V Trifuraceae, p. 1282. 6. Inducing diseases of the mumps group. Family VI. Rabulaceae, p. 1284. FAMILY I. BORRELINACEAE FAM. NOV. Viruses causing polyhedral, wilt, and other diseases in arthropods. The genus Bor- relina Paillot was originally spelled Borrellina by error; from Borrel, name of French scientist. Key to the genera of family Borrelinaceae. I. Known only as attacking lepidopterous insects. Genus I. Borrelina, p. 1225. II. Known only as attacking the honey bee, a hymenopterous insect. Genus II. Morator. p. 1227. Genus I. Borrelina Paillot. (Compt. rend. Acad. Sci., Paris, 182, 1926, 182.) Viruses inducing polyhedral, wilt, and other diseases; hosts, Lepidoptera, so far as known. The type species is Borrelina bombycis Paillot. Key to the species of genus Borrelina. I. Attacking silkworm. II. Attacking nun moth. III. Attacking gypsy moth. IV. Attacking cabbage worm. 1. Borrelina bombycis. 2. Borrelina efficiens. 3. Borrelina reprimens. 4. Borrelina brassicae. 5. Borrelina pieris. 1226 MANUAL OF DETERMINATIVE BACTERIOLOGY 1. Borrelina bombycis Paillot. (Compt. rend. Acad. Sci., Paris, 182, 1926, 182.) Frona Latin bombyx, silk- worm. (Note : Coccus-like bodies sur- rounded by non-staining substances, as- sociated with the induced disease, re- ceived the provisional name Chlamy- dozoon bombycis from Prowazek, Arch. f. Protistenkunde, 10, 1907, 363.) Common names : Silkworm-jaundice virus, silkworm-grasserie virus, silkworm \nlt virus, Gelbsucht virus, Fettsucht \irus. Host: BOMB YCI DAE— Bombyx mori (L.), silkworm. Geographical distribution: Japan, Italy, France. Induced disease : In silkworm, after prodromal period of 5 days or more, yel- low spots on skin, polyhedral bodies in blood, inactivity, loss of appetite, ir- ritability, weakening of body facilitating rupture from mechanical stress, eventual death. Transmission : By feeding. Experi - mentally, also by injection. Serological relationships : Specific ag- glutination, precipitation, and comple- ment fixation. Thermal inactivation : At 60° C in 15 to 20 minutes in blood. Filterability : Passes Berkefeld N and V, Chamberland Li, Lo, and L3 filters. Other properties : May survive at least 2 years in dry state. Stable between pH 5 and about pH 9. Sedimentation constant 17 S. Literature: Aoki and Chigasaki, Cent, f. Bakt., I Abt., Orig., 86, 1921, 481-485; Glaser and Lacaillade, Am. Jour. Hyg., 20, 1934, 454-464; Glaser and Stanley, Jour. Exp. Med., 77, 1943, 451-466; v. Prowazek, Cent. f. Bakt., I Abt., Orig., 67, 1912, 268-284; Suzuki, Bull. Imperial Kyoto Sericultural College, 1, 1929, 45- 75; Trager, Jour. Exp. Med., 61, 1935, 501-513. 2. Borrelina eflGiciens spec. nov. From Latin efficiens, effective, in reference to effectiveness of this virus in controlling nun-moth infestations. Common names : Nun-moth disease virus, nun-moth wilt virus, Wipfelkrank- heit virus. Host: LYMANTRIIDAE—Lyman- tria monacha (L.), nun moth. Geographical distribution: Europe. Induced disease : In eggs, larvae, pupae and occasionally adults of nun moth, polyhedral bodies in affected tissues. Blood of sick larvae turbid; later, blood cells few ; contents of body finally become a gray-brown, semifluid mass. Transmission : By feeding. Thermal inactivation : At 55 to 60° C in 5 to 10 minutes. Filterability : Fails to pass Berkefeld and Chamberland filters. Other properties : May remain viable at least 2 years in dry state. Literature : Escherich and Miyajima, Naturwissens. Ztschr. f. Forst- u. Land- wirtschaft, 9, 1911, 381-402; Wachtl and Kornauth, Mitth. a. d. forstl. Versuch- swesen Osterreichs, 16, 1893, 1-38; Wahl, Centralbl. Gesam. Forstw., 35, 1909, 164- 172; 212-215; 36, 1910, 377-397; 37, 1911, 247-268; 38, 1912, 355-378. 3. Borrelina reprimens spec. nov. F'rom Latin rcprimere, to restrain. Common name : Gypsy-moth wilt virus. Host: LYMANTRIIDAE—Porthe- iria dispar (L.), gypsy moth. Geographical distribution : United States. Induced disease : In gypsy moth cater- pillar, flaccidity, disintegration of tissues, eventual collapse as a watery sack. Death occurs in 13 to 29 (average 21 ) days after infection; caterpillar may remain attached to its support by prolegs; skin ruptures easily. Polyhedral bodies orig- inate in nuclei of the tracheal matrix, hypodermal, fat, and blood cells. Transmission : By feeding on contam- FAMILY BORRELINACEAE 1227 inated foliage. Not through undamaged skin. Filterability : Passes Berkefeld N, not Pasteur-Chamberland F, filter. Literature : Chapman and Glaser, Jour. Econ. Entomol., 8, 1915, 140-150; 9, 1916, 149-167; Glaser, Jour. Agr. Res., 4, 1915, 101-128; Science, 4S, 1918, 301-302; Glaser and Chapman, Jour. Econ. En- tomol., 6, 1913, 479-488. 4. Borrelina brassicae Paillot. (Compt. rend. Acad. Sci., Paris, 182, 1926, 182.) From name of host, Pier is brassicae. Common name : Cabbage -worm grass - erie virus. Host : PIERIDAE — Pieris brassicae (L.), cabbage worm. Induced disease : In cabbage worm, no nuclear or cytoplasmic inclusions; nuclei of fat and hj^podermal cells hypertrophied and soon disorganized. Transmission : By feeding. Other properties: Described as sub- microscopic in size, intracytoplasmic. Literature : Paillot, loc. cit. ; Ann. Inst. Pasteur, 40, 1926, 314-452; L'infection chez les insectes. Immunite et sym- biose, 535 pages, Trevoux, Patissier, 1933. 5. Borrelina pieris Paillot. (Compt. rend. Acad. Sci., Paris, 182, 1926, 182.) From New Latin Pieris, generic name of host. Common name : Virus of nuclear disease of pierids. Host : PIERIDAE — Pieris brassicae (L.), cabbage worm. Induced disease : In cabbage worm, body yellowish below, tears easily just before death; chromatin of nuclei in fat and blood cells condensed in irregular masses; cytoplasmic inclusions staining faintly red in Giemsa preparations. Transmission : By feeding. Other properties : Described as intra- cytoplasmic, less than 0.1 micron in di- ameter. Literature : Paillot, loc. cit. ; Ann. Inst. Pasteur, 40, 1926, 314-452; L'infection chez les insectes. Immunite et sym- biose, 535 pages, Trevoux, Patissier, 1933 . Appendix: Borrelina flacheriae quoted from Paillot, L'infection chez les insects. 535 pp., Trevoux, Patissier, 1935, see p. 96. Cause of gattine in the silkworm, Bombyx wori L. No previous reference to a description of this species has been found. Genus II. Morator gen. nov. Only one species at present, inducing the disease known as sacbrood of the honey bee. Generic name from Latin vioraior, loiterer. The type, and onlj', species is Morator aetatulae spec. nov. 1. Morator aetatulae spec. nov. From Latin actatula, early period of life, in reference to attack on immature stages of host, exclusively. Common name : Honej'-bee sacbrood virus. Host: APIDAE — Apis mellifera L., honey bee (immature stages onljO- Insusceptible species : LYMANTRII- DAE — Porthetria dispar (L.), gypsy moth. Geographical distribution: United States. Induced disease : In the honey bee, immature stages only are susceptible ; in- fected larvae die, usually after capping, some of the dead brood being uncapped by the bees. Occasionally caps are punc- tured. Affected areas of comb are usu- ally small and scattered. Each larva is extended along its cell, head turned up- ward toward the roof. A larva recently dead appears light yellow, light gray, or light brown, soon darkening to brown or almost black. Cuticle of dead larva tough, permitting extraction of the sac- 1228 MANUAL OF DETERMINATIVE BACTERIOLOGY like mass without rupture; contents watery with many suspended, fine, brown particles. There are no characteristic intracellular bodies in aiTected tissues. Dead larvae eventually dry down to form scales that are black and roughened, that separate readily from the cell wall, and that may be lifted out intact. Colonies tend to lose virus spontaneously. Transmission : By contamination of food. Not by hands, clothing, or tools. Perhaps through water supply of insects. Thermal inactival ion : In water, at 58° C in 10 mimites. In honey, at 70 to 73° C in 10 minutes. Filterability : Passes Berkefeld and Pasteur-Chamberland filters. Other properties : Withstands drying 20, not 22, days, exposure to sunlight 7 hours or less, storage in honey a month or more, ^ to 2 per cent aqueous solutions of carbolic acid 3 weeks or more. Literature : McCray and White, U. S. Dept. Agr., Dept. Bull. 671, 1918; White, U. S. Dept. Agr., Bur. of Entomol., Circ. 169, 1913; U. S. Dept. Agr., Dept. Bull. 92, 1914; ihid., Dept. Bull. 431,1917. FAMILY BORRELIOTACEAE 1229 FAMILY II. BORRELIOTACEAE FAM. NOV. Viruses of the Pox Group, inducing diseases characterized in general by discrete primary and secondary lesions of the nature of macules, papules, vesicles, or pustules. Key to the genera of family Borreliotaceae. I. ^'iruses of the Typical Pox-Disease Group. Genus I. Borreliota, p. 1229. II. Viruses of the Varicella Group. Genus II. Briar eus, p. 1233. III. Viruses of the Herpes Group. Genus III. Scelus, p. 1234. IV. Viruses of the Foot-and-Mouth-Disease Group. Genus IV. Hostis, p. 1239. V. Viruses of the Wart -Disease Group. Genus V. Molitor, p. 1240, Gemis I. Borreliota Goodpasture. (Science, 77, 1933, 121.) Viruses of the Tj-pical Pox-Disease Group, inducing diseases characterized by forma- tion of papules, pustules, and scabs, shed with or without scarring. Generic name from Barrel, investigator who first discovered the specific granules of fowl pox and Latinized name of the smallest Greek letter, iota, signifying smallest particle. The name Cytoryctes variolae Guarnieri 1892 was based on intracellular inclusions, Guar- nieri bodies, as supposed sporozoan parasites (Calkins, Jour. Med. Res., 11, 1904, 136-172). The type species is Borreliota avium Goodpasture. Key to the species of genus Borreliota. I. Affecting domestic fowl. 1. Borreliota avium. II. Affecting man principally, although strains have become adapted to cow, rabbit, etc. 2. Borreliota variolae. III. Affecting swine. 3. Borreliota suis. 1. Borreliota avium (Lipschiitz) Good- the names Kikuth's canary virus and pasture. (Strongyloplastna avium Lip- pigeon-pox virus. schutz, in Kolle, Kraus and Uhlenhuth, Hosts: Chicken, turkey, pigeon, goose, Handbuch der pathogenen Mikroorgan- duck, guinea fowl, quail, hawk, pheasant, ismen, 3 Auft., 8, 1930, 314; Goodpasture, partridge, bunting sparrow, canary. Ex- Science, 77, 1933, 121.) From Latin perimentally, also English sparrow, chick aves, fowl of the air. embryo. Common names: Fowl-pox virus; also Insusceptible species: Man, goat, known as poultry -pox virus, chicken-pox sheep, mouse, rat, guinea pig. virus (but not the virus of the same name Geographical distribution : Europe, attacking man rather than the chicken), Asia, Xorth America; perhaps coexten- or virus of epithelioma contagiosum of sive with the area in which chickens are fowls; strains have been studied under grown under conditions of domestication. 1230 MANUAL OF DETERMINATIVE BACTERIOLOGY Induced disease : In chicken, hyper- plastic nodular lesions of the skin, diph- theritic membranes in mouth and throat, discharges from eyes and nose ; nodules eventually dry up and fall off, usually without leaving scars. Inclusion bodies, known as Bollinger bodies, believed to represent aggregates of minute Borrel bodies or virus particles, leave much grayish-white ash when incinerated ; break readily after digestion by 1 per cent trypsin in 0.2 per cent sodium bicarbon- ate. Borrel bodies coccoid, 0.25 microns in diameter. On chorioallantoic mem- brane of chick embryo, proliferation and hyperplasia, or necrosis. Transmission: By contact, perhaps through wound infection. By blood- sucking dipterous insects. Experimen- tally, by scarification of skin or buccal mucosa; by intravenous, intradermal, subcutaneous, intramuscular, or intra- peritoneal inoculation. May be passed in series by nasal instillation in chickens, obvious mucosal changes occurring only occasionally. Experimentally, by mos- quitoes (CULICIDAE), Aedes aegypti L., A. slimvlans Walker, A. vexans Meigen (as long as 27 days from time of feeding on infective material), and Cidex pipiens L. (indefinitely after infective feeding, as long as the individual mos- quito lives) ; in C. pipiens, the virus has been found also under natural conditions. Serological relationships : Neutralizing and elementary-body-agglutinating anti- sera specific. Antivaccinial serum from rabbit ineffective against fowl-pox virus, although neutralizing vaccinia virus. Immunological relationships : No cross immunity with respect to vaccinia virus in the chicken. Thermal inactivation : At 60° C in 8 minutes ; at 56° C in 30 minutes. Filterability : Passes Berkefeld V, not Chamberland L2, filter candle. Other properties : Drying at room tem- perature in vacvo does not inactivate. Viable after storage at least 24 months at 0 to 4° C, dry. Strains: A strain known as Kikuth's canary virus has been studied in some detail. When introduced into the rabbit it induces formation of neutralizing anti- bodies that react strongly with homol- ogous virus, moderately against fowl-pox virus. Antivaccinial serum is ineffective against it. In canaries, it induces pro- liferation of dermal epithelium with cytoplasmic inclusions, the inflammatory process being characterized by predom- inantly mononuclear cells with vacuo- lated cytoplasm; in the lung there is massive accumulation of large mononu- clear cells containing the specific cyto- plasmic inclusions; the disease is regu- larly fatal. Passes Berkefeld N filter. Size estimated as 120 millimicrons by centrifugation. (Bechhold and Schles- inger, Ztschr. f. Hyg., 115, 1933, 354- 357; Burnet, Jour. Path, and Bact., 37, 1933, 107-122; Burnet and Lush, Brit. Jour. Exp. Path., 17, 1936, 302-307; Gaede, Cent. f. Bakt., I Abt., Orig., 135, 1935, 342-346; Kikuth and Gollub, ibid., 125, 1932, 313-320.) Literature: Andervont, Am. Jour. Hyg., 6, 1926, 719-754 ; Brandly and Dun- lap, Jour. Am. Vet. Med. Assoc, 95, 1939, 340-349; Brandly et al.. Am. Jour. Vet. Res., 3, 1941, 190-192; Brody, Cornell Agr. Exp. Sta. (Ithaca). Memoir 195, 1936; Buddingh, Jour. Exp. Med., 67, 1938, 933-940; Burnet and Lush, Brit. Jour. Exp. Path., 17, 1936, 302-307; Danks, Am. Jour. Path., 8, 1932, 711-716; Findlay, Proc. Roy. Soc. London, Ser. B, 102, 1928, 354-379; Goodpasture and A. M. Woodruff, Am. Jour. Path., 6, 1930, 699-712; Goodpasture and C. E. Wood- ruff, ibid., 7, 1931, 1-8; Irons, Am. Jour. Hyg., 20, 1934, 329-351 ; Kligler and Ash- ner, Proc. Soc. Exp. Biol, and Med., 28, 1931, 463-465; Kligler et al., Jour. Exp. Med., 49, 1929, 649-660; Ledingham, Lancet, 221, 1931 {2), 525-526; Ludford and Findlay, Brit. Jour. Exp. Path., 7 1926, 256-264; Matheson et al, Poultry Science, 10, 1931, 211-223; Megrail, Am. Jour. Hyg., 9, 1929, 462-465; Nelson, Jour. Exp. Med., 74, 1941, 203-212; A. M. Woodruff, and Goodpasture, Am. Jour. FAMILY BORRELIOTACEAE 1231 Path., 7, 1931, 209-222; C. E. Woodruff, ibid., 6, 1930, 169-174; C. E.Woodruff, and Goodpasture, ibid., 5, 1929, 1-10; 6, 1930, 713-720. 2. Borreliota variolae (Lipschiitz) Goodpasture. (Sirongyloplasma variolae Lipschiitz, in KoUe, Kraus and Uhlen- huth, Handbuch der pathogenen Mikro- organismen, 3 Aufl., 8, 1930, 317; Good- pasture, Science, 77, 1933, 121.) From New Latin variola, smallpox. Common names: Variola virus, small- pox virus. Most studies of this virus have been concerned with the vaccinia strain ; see Strains below. Hosts : Man, cow and rabbit are sus- ceptible to strains that appear especiallj' adapted to them (see Sirai7is below). Experimentally, also chicken (and chick embryo); Chrysemys marginata, turtle; guinea pig, horse, pig; Macaca miilatta (Zimmermann), rhesus monkey; M. irus, cynomolgus monkey; orang-outang; Ma- cacus fuscatus. Geographical distribution: Nearly world-wide, except where excluded bj' isolation or protective vaccination. Induced disease : In man, mild to severe smallpox, sometimes with pocks few and discrete but often with pocks numerous and coalescing; onset sudden, 6 to 22 days (average 12) after infection; headache, vomiting, fever, often rashes on bodj' before appearance of the specific eruption, bright red spots becoming vesic- ular and eventually pustular; the pocks are commonest on face, forearms, wrists, palms of hands, and soles of feet ; pustules gradually become flattened scabs and drop off, leaving no scar if superficial and not secondarily infected; in hemorrhagic smallpox there are numerous hemorrhages into the skin and mortality is high, death often preceding formation of pustules; severity of disease and mortality roughly proportional to the amount of eruption on the face. Transmission : By contact with in- fected individuals or contaminated ar- ticles; perhaps by droplet infection, obvious primary lesions characterizing experimental transmission by scarification but not natural spread. Serological relationships : Hyperim- mune calf serum neutralizes virus. Neu- tralization depends on an antibody not involved in agglutination and precipita- tion. Antivaccinial serum gives com- plement fixation in the presence of variola virus. One agglutinogen (L) labile at 56° C, one (S) stable at 95° C; both are parts of a single protein but can be de- graded independently; chymotrypsin de- stroj's activity of S, not L. Increasing neutralization in immune serum and virus mixtures in vitro with progressive incuba- tion; partial reactivation on simple dilu- tion. Antivaccinial sera agglutinate Pas- chen bodies of vaccinia but not Borrel bodies of fowl pox ; anti-fowl -pox sera ag- glutinate Borrel but not Paschen bodies. No cross reactions with herpes virus. Immunological relationships : In vac- cinia-immune swine, protective sub- stances pass via colostrum, conveying passive immunity to young for 2 to 3 months after birth. In man, immunity against variola virus is conferred by ear- lier infection with vaccinia strain. In hen, previous infection with fowl-pox virus does not immunize with respect to vaccinia virus. Thermal inactivation : At 55° C in 20 minutes. Filterability : Passes Berkefeld V, not Mandler, filter. Other properties: Density about 1.16. Sedimentation constant 5000 X lO^i^ (corrected to water at 20° C). Retains activity in glycerine best at pH 7.0. 0.1 per cent gelatin delays spontaneous in- activation at 5 to 10° C. Withstands absolute alcohol, ether, acetone, and petroleum ether 1 hour in dry samples at 4° C without decrease in activity. Inac- tivated without disruption by sonic vi- brations of about 8900 cycles per second. Diameter estimated as 125 to 175 milli- microns by filtration; 236 to 252 milli- microns by ultracentrifugation. Elec- tron micrographs show limiting surface 1232 MANUAL OF DETERMINATIVE BACTERIOLOGY membrane, dense granules (usually 5) within; tendency to rectangular outlines with rounded corners. At least 5.6 per cent of virus is reported to be thymo- nucleic acid. Contains nitrogen, 15.3 per cent; carbon, 33.7 per cent; phosphorus, 0.57 per cent; phospholipid (lecithin), 2.2 per cent; neutral fat, 2.2 per cent; reducing sugars after hydrolysis, 2.8 per cent; cystine, 1.9 per cent; copper, 0.05 per cent. Strains: Besides the typical variola strain, var. hominis Goodpasture (Sci- ence, 77, 1933, 121), several distinctive strains have been studied. A spontane- ous cowpox strain differs in some antigens but affords cross immunity with respect to var. bovis Goodpasture {loc. cit., 121), vaccinia virus, which in turn immunizes against typical variola virus. A spon- taneous rabbit-pox strain, serologically resembling neurovaccine virus, is be- lieved to exist independently in Europe and the United States. The varieties equi (horse-pox virus), porci (swine strain), and avium (sheep and goat pox virus) have been attributed to this species by Goodpasture {loc. cit., 121). The alastrim strain (causing variola minor) differs from the type in producing a relatively mild disease in man and in inducing the formation of a distinctive type of intracellular inclusion in affected tissues. Literature: Amies, Jour. Path, and Bact., 47, 1938, 205-222; Andervont, Am. Jour. Hyg., 7, 1927, 804-810; Behrens and Ferguson, Jour. Inf. Dis., 56, 1935, 84-88; Behrens and Nielson, ibid., 56, 1935, 41-48; Buddingh, Am. Jour. Hyg., 38, 1943, 310-322; Craigie and Wishart, Brit. Jour. Exp. Path., 15, 1934, 390-398 ; Jour. Exp. Med., 64, 1936, 819-830; Bearing, Am. Jour. Hyg., 20, 1934, 432-443; Doug- las et al.. Jour. Path, and Bact., 82, 1929, 99-120; Downie, Brit. Jour. Exp. Path., 20, 1939, 158-176; Eagles, ibid., 16, 1935, 181-188; Elford and Andre wes, Brit. Jour. Exp. Path., 13, 1932, 36-42; Good- pasture, Woodruff, and Buddingh, Am. Jour. Path., 8, 1932, 271-282 ; Green et al., Jour. Exp. Med., 75, 1942, 651-656, Greene, ibid., 61, 1935, 807-831 ; Herzberg, Ztschr. Immunitatsforsch. u. exper. Therap., 86, 1935, 417-441; Hoagland et al., Jour. Exp. Med., 71, 1940, 737-750; 72, 1940, 139-147; 7 4, 1941, 69-80, 133-144; 76, 1942, 163-173; Hu et al.. Jour. E.xp. Med., 63, 1936, 353-378; Keogh, Jour. Path, and Bact., ^, 1936, 441-454; Ledingham, Brit. Jour. Exp. Path., 5, 1924, 332-349; Jour. Path, and Bact., 35, 1932, 140-142; Macfarlane and Dolby, Brit. Jour. Exp. Path., 21, 1940, 219-227; Macfarlane and Salaman, ibid., 19, 1938, 184-191; McFarlane et al., ibid., 20, 1939, 485-501 ; Moriyama, Arch, f . Virus- forsch., 1, 1940, 422-429; Nelson, Jour. Exp. Med., 60, 1934, 287-291; 78, 1943, 231-239; Nye and Parker, Am. Jour. Path., 5, 1929, 147-155; Parker, Jour. Exp. Med., 67, 1938, 361-367, 725-738; Parker and Muckenfuss, Jour. Infect. Dis., 53, 1933, 44-54; Parker and Rivers, Jour. Exp. Med., 62, 1935, 65-72; 64, 1936, 439-452; 65, 1937, 243-249; Pasc hen, Deutsch. med. Wchschr., 89, 1913, 2132- 2136; Pearce et al., Jour. Exp. Med., 68, 1936, 241-258, 491-507; Jour. Path, and Bact., 43, 1936, 299-312; Pickels and Smadel, Jour. Exp. Med., 68, 1938, 583- 606; Rhodes and van Rooyen, Jour. Path, and Bact., 44, 1937, 357-363; Rivers and Ward, Jour. Exp. Med., 58, 1933, 635-648; 62, 1935, 549-560; Rivers et al., ibid., 65, 1937, 677-685; 69, 1939, 857-866; Rosahn et al.. Jour. Exp. Med., 63, 1936, 259-276, 379-396; Rosenau and Andervont, Am. Jour. Hyg., 13, 1931, 728-740; Salaman, Brit. Jour. Exp. Path., 18, 1937, 245-258; Shedlovsky and Smadel, Jour. Exp. Med., 75, 1942, 165-178; Smadel and Rivers, ibid., 75, 1942, 151-164; Smadel et al., ibid., 68, 1938, 607-627; 7i, 1940, 373-389; 77, 1943, 165-171 ; Smith, Jour. Path, and Bact., 88, 1930, 273-282; Sprunt, Proc. Soc. E.xp. Biol, and Med., 51, 1942, 226- 227; Jour. Exp. Med., 75, 1942, 297-304; Stritar and Hudson, Am. Jour. Path., 12, 1936, 165-174; Ward, Jour. Exp. Med., 50, 1929, 31-40. 3. Borreliota suis spec. nov. From Latin sus, swine. Common name: Swine-pox virus (not FAMILY BORRELIOTACEAE 1233 the vaccinia strain of variola virus in swine). Host : S UIDAE—Sus scrofa L., domes- tic swine. Insusceptible species : Rabbit. Geographical distribution: United States (Iowa). Induced disease : In swine, locally, red- dened hyperemic papules 3 to 7 mm in diameter; papules become briefly vesicu- lar, then change gradually to true pus- tules, finally forming dark brown to blackish scabs which are shed after a few weeks without scarring; no secondary lesions in hogs free from lice, but in in- fested animals numerous secondary le- sions appear 1 to 2 weeks after primar}' lesions and are commonly most numerous in the inguinal and axillary regions. Mortality negligible but growth retarded. Virus has been recovered from hog louse after feeding on affected swine. Transmission : By hog louse, Haemato- pinus suis (HAEMATOPINIDAE), probably mechanically. B\' experi- mental scarification of skin. Serological relationships : Xo reaction with neutralizing sera specific for vac- cinia virus. Immunological relationships: Specific: immunitjf in swine after attack, but no cross immunity with respect to vaccinia virus. Filterability : Passes Berkefeld V and N filters. Literature : Csontos and von Xyiredy, Deutsch. tierarztl. Wchnschr., 41, 19.33, 529-532; Schwarte and Biester, Am. Jour. Vet. Res., 2, 1941, 13G-140; Shope, Arch, f. Virusforsch., 1, 1940, 4.57-467. Genus II. Briareus ge7i. nov. Viruses of the Varicella Group, causing diseases characterized by reddened spots and rings in affected tissues, becoming papular or vesicular. Generic name from Latin Briareus, name of a hundred-armed giant. The t5^pe species is Briareus varicellae spec. nov. Key to the species of genus Briareus. I. Causing chicken pox and herpes zoster in man. II. Causing measles in man. 1. Briareus varicellae spec. nov. From New Latin varicella, chicken pox. Common names : Varicella virus, chic- ken-pox virus; much evidence for iden- tity with so-called herpes-zoster virus has been presented. Host : HOMIN IDAS— Homo sapiens L., man. Insusceptible species: Chick embryo. Geographical distribution: World-wide. Induced disease : In man, usually abrupt onset, rash at first macular, soon papular and vesicular; vesicles generally discrete, soon rupturing, healing with scab formation and itching ; separation of deeper scabs may leave persistent scars ; in severe cases there may be stomatitis, laryngitis, and nasal lesions. In human skin grafted on chorioallantois of chick 1. Briareus varicellae. 2. Briareus morhillorum. embryo, experimentally, pustular lesions as in natural disease, with intranuclear acidophilic inclusions ; no gross vesicula- tion. Transmission: By contact. Bj' spread of droplets. Children in contact with herpes zoster patients sometimes con- tract varicella. Serological relationships : Majority of herpes zoster sera that agglutinate zoster antigen also agglutinate elementary' bodies of varicella; complement fixation tests also indicate relationship of virus from herpes zoster and varicella. Chic- ken-pox sera do not flocculate smallpox brain-virus antigen. Immunological relationships : Children previously having varicella are immune to inoculation with herpes zoster virus. 1234 MANUAL OF DP]TERMINATIVE BACTERIOLOGY Literature : Amies, Brit. Jour. Exp. Path., 15, 1934,314-320; Brain, ibid., U, 1933, 67-73; Bruusgaard, Brit. Jour. Derm. Syph., 44, 1932, 1-24 ; Goodpasture and Anderson, Am. Jour. Path., 20, 1944, 447-453; Havens and Mayfield, Jour. Inf. Dis., 50, 1932, 242-248; Irons et al.. Am. Jour. Hyg., 33, (B), 1941, 50-55; Kundratitz, Monatsschr. Kinderheilk., 29, 1925, 516-523; Lipschiitz and Kundra- titz, Wien. klin. Woch., 38, 1925, 499-503. 2. Briareus morbillorum spec. nov. From New Latin ynorhilli, measles. Common name : Measles virus. Host: HOMINIDAE—Honw sapiens L., man. Experimentally, also CERCO- PITHEC ID AE— Macaco, mulalta (Zim- mermann), rhesus monkej'. P H ASI- AN I DAE— Gallus gallus (L.), chick embryo (no lesions, but 30 serial pas- sages). Geographical distribution: World-wide except in isolated communities. Induced disease : In man, after incuba- tion period of 7 to 21 days, bright red spots on buccal mucosa, especially near first molar tooth (Koplik's spots) followed by rash on face, head, neck, then arms. trunk, and legs; papules often crescents, lesions usually discrete ; rash fades, leav- ing brownish discoloration and desqua- mation. Transmission: By contact. By drop- lets. Serological relationships : Convalescent serum is reported to modify the course of the induced disease if administered in- travenously in the preeruptive stage. Immunological relationships : Specific immunity in man after attack. Thermal inactivation : At 55° C in 15 minutes. Filterability : Passes Berkefeld N filter candle and Seitz EK disks. Other properties: Viable at —35° C for at least 4 weeks. Not inactivated by 10 per cent anesthetic ether in 40 minutes. Literature : Blake and Trask, Jour. Exp. Med., 33, 1921, 385-412; Gordon and Knighton, Am. Jour. Path., 17, 1941, 165-176; Hedrich, Am. Jour. Hyg., 17, 1933, 613-636 ; Kohn et al., Jour. Am. Med. Assoc, 111, 1938, 2361-2364; Rake and Shaffer, Jour. Immunol., 38, 1940, 177-200; Kakeet al.. Jour Inf. Dis.,eS, 1941,65-69; Scott and Simon, Am. Jour. Hyg., 5, 1925, 109-126. Genus III. Scelus gen. nov. Viruses of the Herpes Group, inducing diseases characterized in general by vesicular primary lesions, sometimes with subsequent involvement of the nervous system. Generic name from Latin scelus, rascal. The type species is Scelus recurrens spec. nov. Key to the species of genus Scelus. I. In man, cause of so-called fever blisters, herpes febrilis. 1. Scelus recurrens . II. In swine, cause of pseudorabies. 2. Scelus suillum. III. In monkey. 3. Scelus beta. IV. In rabbit, course of the induced disease in nature unknown. 4. Scelus teriium. V. In sheep, cause of ovine balano-posthitis. 5. Scelus ulcer is. VI. In mice, cause of ectromelia. 6. Scelus marmorans. VII. In cattle, cause of erosive stomatitis. 7. Scelus bovinum. FAMILY BORRELIOTACEAE 1235 1. Scelus recurrens spec. nov. From Latin recnrrere, to recur. Note : The name A'eurocystis herpetii Levaditi and Schoen (Compt. rend. Soc. Biol., Paris, 96, 1927, 961 ) was applied provisionally to the causative microorganism of herpes, in the expectation that future research would show inclusion bodies in affected tissues to be stages in its life cycle. Common names : Herpes virus, virus of herpes simplex, virus of herpes febrilis (not herpes zoster virus, for which see varicella virus), virus of keratitis dendri- tica, virus of aphthous stomatitis (of man). Host: HOMINIDAE—Homo sapiens L., man. Experimentally, also rabbit, guinea pig, white mouse, cat, goose, hedge- hog, and, though difficult to infect, dog and pigeon. Chick embryo (but not chicken). Also CERCOPITHECIDAE — Cercocebus fuligiriosus E. Geoffrey, Macacus cynomolgus. CEBIDAE — Ce- bus olivaceus. Insusceptible species: White rat; Bufo viridis; Cercopithecus callithrix; chicken (except embryo). Geographical distribution: Probably world-wide. Induced disease : In man, usually acquired in ffrst three years of life, some- times as aphthous stomatitis; virus prob- ably retained often through life, some- times with periodic reappearance of der- mal lesions, which are vesicular and heal soon. In white mouse, by experimental inoculation of skin, small inflamed vesicu- lar primary lesions about 5 days after in- oculation, usually forming scabs and healing a few days later, but sometimes persisting; if on tail, followed by swelling and paralysis of tail, ascending paralysis and death, or by recovery with acquired immunity; if near head, followed by en- cephalitis and death ; intraperitoneal and sometimes other inoculations immunize ; relapse with recurrence of primary lesions rare. In chick embryo, white, opaque, circular or ring-like primary lesions of small size on chorioallantoic membrane, with or without necrotic secondary lesions in liver, heart, lungs, spleen, and kidneys ; virus enters membrane 1 to 4 hours after it is dropped on its surface; primary lesions may be counted in 48 hours. Transmission: Bj- contacts. Experi- mentally, by skin scarification ; in guinea pig, by feeding. Serological relationships : Distant rela- tionship to pseudorabies virus, Scelus suillum, shown by moderate protection against this virus conferred by some anti- herpes sera. No relationship to vaccinia virus or to virus III of rabbits demon- strable by neutralization tests. Specific complement fixation. Neutralizing anti- body forms reversible union with virus, at least for a time, though with strong mixtures partial irreversibility finally occurs. Immunological relationships: Formal- inized virus and non-lethal strains of virus immunize specifically. No cross immunity with vaccinia virus. Thermal inactivation : At 50 to 52° C in 30 minutes, when moist; at 90 to 100° C in 30 minutes, when dry. At 41.5° C in 50 to 80 hours. Filterability : Passes Berkefeld V filter with slight loss. Other properties : Diameter, by centrif- ugation, computed as 180 to 220 milli- microns; by filtration, 100 to 150 millimicrons. Specific gravity, 1.15. In- activated by repeated freezing and thaw- ing; also by pressure of 3000 atmospheres for 30 minutes. Viable dry at least 18 months at 4° C, in 50 per cent glycerine at least 6 months. Not inactivated at 4° C in 1 per cent aqueous gentian violet. Charged negatively in solutions of hy- drogen-ion concentration up to about pH 8. Isoelectric point, pH 7.2 to 7.6. In- activated by incubation in vitro at pH 6 with synthetic vitamin C (ascorbic acid). Literature : Anderson, Science, 90, 1939, 497; Am. Jour. Path., 16, 1940, 137- 156; Andervont, Jour. Inf. Dis., U, 1929, 383-393; 45, 1929, 366-385; 49, 1931, 507- 529; Andrewes, Jour. Path, and Bact., 33, 1930, 301-312; Bassett et al., Compt. rend. Acad. Sci., Paris, 200, 1935, 1882- 1884; Bechhold and Schlesinger, Ztschr. 123G MANUAL OF DETERMINATIVE BACTERIOLOGY f. Hyg., 115, 1933, 342-353 ; Be dsoii, Brit. Jour. Exp. Path., 12, 1931, 254-260; Bedson and Bland, ibid., 9, 1928, 174-178; Blanc and Caminopetros, Compt. rend. Soc. Biol., Paris, 84, 1921, 859-860; Boak et al., Jour. Exp. Med., 71, 1940, 169-173; Brain, Brit. Jour. Exp. Path., IS, 1932, 166-171 ; Buggs and Green, Jour. Inf. Dis., 58, 1936, 98-104; Burnet and Lush, Jour. Path, and Bact., 48, 1939, 275-286; 49, 1939, 241-259; Lancet, 236, 1939 (1), 629-631; Burnet et al., Austral. Jour. Exp. Biol, and Med. Sci., 17, 1939, 35-40; Dawson, Am. Jour. Path., 9, 1933, 1-6; Elford et al.. Jour. Path, and Bact., 86, 1933, 49-54; Findlay and MacCallum, Lancet, 238, 1940 {1), 259-261 ; Fischl and Schaefer, Klin. Wochnschr., 8, 1929, 2139-2143; Flexner, Jour. Gen. Physiol., 8, 1927, 713-726; Jour. Exp. Med., 47, 1928, 9-36; Friedenwald, Arch. Ophthal- mol., 52, 1923, 105-131; Goodpasture, Medicine, 8, 1929, 223-243; Goodpasture and Teague, Jour. Med. Res., 44, 1923, 121-138; Gunderson, Arch. OpthalmoL, 15, 1936, 225-249 ; Holden, Jour. Lif. Dis., 50, 1932, 218-236; Keddie and Epstein, Jour. Am. Med. Assoc, 117, 1941, 1327- 1330; Levaditi and Lepine, Compt. rend. Acad. Sci., Paris, 189, 1929, 66-68; Leva- diti and Nicolau, Compt. rend. Soc. Biol., Paris, 90, 1924, 1372-1375; Long, Jour. Clin. Investigation, 12, 1933, 1119-1125; Magrassi, Boll. 1st. Sieroterap. Milanese, 14, 1935, 773-790; McKinley, Proc. Soc. Exp. Biol, and Med., 26, 1928, 21-22; Naegeli and Zurukzoglu, Cent. f. Bakt. I Abt., Orig., 135, 1935, 297-299 ; Nicolau and Kopciowska, Ann. Inst. Pasteur, 60, 1938, 401-431 ; Parker and Nye, Am. Jour. Path., 1, 1925, 337-340; Perdrau, Proc. Roy. Soc. London, Ser. B, 109, 1931, 304- 308; Jour. Path, and Bact., 47, 1938, 447- 455; Remlinger and Bailly, Comp. rend. Soc. Biol., Paris, 94, 1926, 734-736; 1064- 1066; 95, 1926, 1542-1545; 96, 1927, 404- 406; 1126-1128; 97, 1927, 109-111; Sabin, Brit. Jour. Exp. Path., 15, 1934, 372-380; Schultz and Hoyt, Jour. Immunol., 15, 1928, 411-419; Shaffer and Enders, ibid., 37, 1939, 383-411; Simon, International Clinics, Series 37, 3, 1927, 123-128; Smith et al.. Am. Jour, Path., 17, 1941, 55-68 Warren et al.. Jour. Exp. Med., 71, 1940 155-168; Weyer, Proc. Soc. Exp. Biol and Med., 30, 1932, 309-313; Zinsser Jour. Exp. Med., 49, 1929, 661-670 Zinsser and Seastone, Jour. Immunol. 18, 1930, 1-9; Zurukzoglu and Hruszek Cent. f. Bakt. I Abt., Orig., 128, 1933 1-12. 2. Scelus suillum spec. nov. From Latin siiillus, pertaining to swine. Common names: Pseudorabies virus, mad-itch virus. Hosts : Domestic cattle, swine, dog, cat, horse, sheep. Experimentally, also rab- bit, guinea pig, white rat, white mouse, gray field mouse, duck, chicken, chick embryo; Macaca mvlaita (Zimmermann), rhesus monkey. Geographical distribution: France, Germany, Hungary, Holland, Denmark, Switzerland, Siberia, Brazil, United States. Induced disease : In cattle, licking of affected area, usually somewhere on hind- quarters, sudden decrease in milk pro- duction in dairy animals, violent rubbing, biting, and gnawing of lesion; swelling and discoloration of affected parts with oozing of serosanguineous fluid ; grinding of teeth and excessive salivation in some individuals; death, preceded by clonic convulsions, violent tossing of head, and shallow respiration, usually 36 to 48 hours after onset. In pig, mild but highly con- tagious disease ; slight nerve-cell degen- eration, predominance of vascular and interstitial lesions. Transmission : By contact in swine, not in cattle. By feeding in cats, brown rats, and swine. Serological relationships : Cross neu- tralization between constituent strains. Anti-herpes sera protect in some cases against small, but constantly infective, doses of pseudorabies virus. Literature: Aujeszky, Cent. f. Bakt. I Abt., Orig., 32, 1902, 353-357; F. B. Bang, Jour. Exp. Med., 76, 1942, 263-270; FAMILY BORRELIOTACEAE 1237 O. Bang, Acta path, et inicrobiol. Scand., Suppl., 11, 1932, 180-182; Carini and Ma- ciel, Bull. Soc. Path, exot., 5, 1912, 576- 578; F0lger, Acta path, et microbiol. Scand., Suppl., 11, 1932, 182-187; Glover, Brit. Jour. Exp. Path., 20, 1939, 150-158; Gowen and Schott, Am. Jour. Hj^g., 18, 1933, 674-687; Hurst, Jour. Exp. Med., 58, 1933, 415-433; 69, 1934, 729-749; 63, 1936, 449-463; Koves and Hirt, Arch, wissensch. u. prakt. Tierheilk., 68, 1934, 1-23; Morrill and Graham, Am. Jour. Vet. Res., 2, 1941, 35-40; Brit. Jour. E.xp. Path., 15, 1934, 372-380; Shope, Proc. Soc. Exp. Biol, and Med., 30, 1932, 308- 309; Jour. Exp. Med., 5J!^, 1931, 233-248; 57, 1933, 925-931 ; 62, 1935, 85-99, 101-117, Traub, ihid., 58, 1933, 663-681; 61, 1935, 833-838. 3. Scelus beta spec. nov. From beta, second letter of Greek alphabet, in ref- erence to common name. Common name : B virus. Hosts : HOMINIDAE — Homo sapiens L., man. CERCOPITHECIDAE—Ma- caca mulatta (Zimmermann), rhesus monkey. E.xperimentally, also LEPOR- IDAE — Oryciolagus cuniculus (L.), rab- bit. CAVIIDAE—Cavia-porcellus (L.), guinea pig. Geographical distribution: United States (from captive monkeys and man). Induced disease : In man, local and rela- tively insignificant lesion on bitten part, later flaccid paralysis of legs, urinary retention, ascending paralysis, and death by respiratory failure. In Macaca viu- latta, experimentally by intracutaneous injection, hemorrhagic or vesiculo-pustu- lar lesions without later involvement of central nervous system but with subse- quent acquired immunity. Acidophilic intranuclear inclusions in lesions. Transmission : To man, by bite of mon- key. To monkey, experimentallj', by injection. Literature : Burnet et al., Austral. Jour. Exp. Biol, and Med. Sci., 17, 1939, 35-40, 41-51; Sabin, Brit. Jour. Exp. Path., 15, 1934, 248-268, 268-279, 321-334 ; Sabin and Hurst, ibid., 16, 1935, 133-148; Sabin and Wright, Jour. Exp. Med., 59, 1934, 115- 136. 4. Scelus tertium spec. nov. From Latin tertius, third. Common name: Virus III of rabbits. Host : LEPORIDAE-^Oryctolagvs cu- niculus (L.), domestic rabbit. Insusceptible species : No obvious dis- ease in inoculated guinea pig, white mouse, monkej^ {Macaca mulatta Zim- mermann), rat, or man; hence the as- sumption that these are naturally im- mune, but they may be merely tolerant or klendusic. Geographical distribution : L'nited States (apparently spontaneous in some individuals of the laboratory rabbit). Induced disease : In domestic rabbit, experimentally, after incubation period of 4 to 6 days, failure to eat, loss of weight , occasionallj^ diarrhea and temperatures of 104 to 107° F; small, superficial, red spots and papules on skin at site of inocu- lation; local infiltration of tissues with endothelial leucocytes, swelling of in- volved epithelial cells ; nuclear inclusions present in endothelial leucocytes and some other cells ; disease not fatal ; virus in circulating blood only during early stages; recovery in a few days without scar formation but with development of specific immunity. The course of the natural disease, presumed to occur in rabbits, is still unknown. Transmission : Experimentally, by in- jection of filtrates from diseased tissues; on several occasions also from blood or tissues of apparently normal rabbits. Serological relationships : Specific neu- tralizing substances occur in the serum of recovered rabbits. Immunological relationships : Specific immunity but no cross reactions with vaccinia or herpes viruses. Thermal inactivation : In 10 minutes at 55° C, but not in 30 minutes at 45° C. Filterability : Passes Berkefeld V and N filters; passes L2 filter candle. Other properties : Viable at least 6 1238 MANUAL OF DETERMINATIVE BACTERIOLOGY weeks in 50 per cent glycerine and 16 months dried when frozen, and stored on ice. Literature : Andrewes, Brit. Jour. Exp. Path., 10, 1929, 188-190, 273-280; Jour. Path, and Pact., 33, 1930, .301-312; 50, 1940, 227-234; Rivers and Stewart, Jour. Exp. Med., 48, 1928, 603-613; Rivers and Tillett, ibid., 39, 1924, 777-802; 40, 1924, 281-287; Topacio and Hyde, Am. Jour. Hyg., 15, 1932, 99-124. 5. Scelus ulceris spec. nov. From Latin ulcus, sore spot. Common name : Ovine balano-posthitis virus. Host: BOVIDAE—Ovis arics L., sheep. Geographical distribution : United States, Australia. Induced disease : In sheep, ulceration with scab production : lesions most severe on prepuce and vulva; in the male, the penis may be involved, usually only with mild inflammation, but if accompanied by parapliimosis there may be extensive ul- ceration and heavy scab formation. Transmission : Venereally. Experi- mentally, by inoculation of prepuce. Filterability : Passes Berkefeld N and W filters, a 7 lb Mandler candle, and a 31 per cent collodion membrane. Literature: Tunnicliff and Matisheck, Science, 94, 1941, 283-284. 6. Scelus marmorans spec. nov. From Latin marmorarc, to marble, in reference to mottling of spleen and liver in host. Common name : Ectromelia virus. Hosts: MURIDAE — Mus miisculus L., white mouse. Esperimentally, also MURIDAE — Rattus norvegicits (Ber- kenhout), rat (infection inapparent). Also, PHASIANIDAE—Gallus galbis (L.), chick embrj^o (12-day-old White Leghorn chick embryo at 36 to 37° (' ; less satisfactory results at higher tem- peratures of incubation or in embryos in spring eggs). Derived strains of this virus infect rabbit and guinea pig, not susceptible to original virus from mouse. Geographical distribution: England. Induced disease: In white mouse, spleen mottled, liver edges translucent, peritoneal fluid increased in amount ; loss of weight; later, cutaneous lesions on foot or elsewhere; affected foot swells, becomes moist, scabbed, then recovers or dries up and separates from the skin at limit of original swelling; in acute dis- ease, death without gross lesions, or, at autopsy, gut dark red, liver dirty gray, soft, bloodless, sometimes mottled, spleen necrotic ; inclusion bodies most numerous in lesions of the skin, round or oval, 4 to 13 microns long, without internal differ- entiation; very young mice probably be- come infected without developing ap- parent disease and remain carriers for some time. In rat, inapparent infection ; after initial increase of virus, circulating antibodies appear and immunity to rein- fection is established. Transmission: In mouse, by contact. In rat, experimentally, bj- intranasal inoculation. Serological relationships: Neutralizing antibodies occur in convalescent mouse serum. Immune sera from the guinea pig specifically agglutinate elementary bodies obtained from infected skin of the white mouse. Immunological relationships : Recov- ered mice are solidly immune to many lethal doses. Thermal inactivation : At 55° C in 30, not in 10, minutes. Filterability : In broth, passes Mandler, Pasteur-Chamberland Lo, and Berkefeld N filters. Other properties: Survives drying 6 months, freezing (—10° C) 2 months, 50 per cent glycerine 5 months at least. Resists 1 per cent phenol 20, not 40, days. Size, estimated by filtration, 100 to 150 millimicrons; by ultraviolet-light photog- raphy, 130 to 140 millimicrons. Literature : Barnard and Elford, Proc. Roy. Soc. London, Ser. B, 109, 1931, 360- 380; Baumgartner, Cent. f. Bakt., I Abt., Orig., 133, 1935, 282-289 ; Burnet and Lush, Jour. Path, and Bact., 42, 1936, 469-476; 43, 1936, 105-120; Jahn, Arch. f. Virus- forsch., 1, 1939, 91-103; Kikuth and FAMILY BORKELIOTACEAE 1239 Gonnert, Arch. f. Virusforsch., 1, 1940, 295-312; Marchal. Jour. Path, and Bact., 3S, 1930, 713-728; McGaughey and White- head, ihid., 37, 1933, 253-256; Paschen, Cent. f. Bakt., I Abt., Orig., 135, 1936, 445-452. 7. Scelus bovinum spec. nov. From Latin borinus, of ox, bull, or cow. Common name : Ercsive-stomatitis virus. Host: BOVIDAE—Bos taurus L., domestic cattle. Experimentally^ also chorioallantoic membrane of developing hen's egg. Insusceptible species: CAVIIDAE — Cavia -porcellus (L.), guinea pig. (In rats, rabbits, mice, sheep, no reaction has been noted after inoculation.) Geographical distribution: South Africa (Natal) ; perhaps Ireland (Armagh -disease virus). Induced disease : In young domestic cattle, lesions on tongue, dental pad, and lips pearl-like at first, then breaking down to form superficial erosions, with white glistening base and red border. Lesions may coalesce to form large, ragged, eroded areas, healing uneventfully with scar formation. No foot lesions ; no excessive salivation; no "hotness" of mouth; no systemic disturbances. Transmission : Spreads slowly, mainly to animals less than three years old, prob- ably by contact. Experimentally, by injection into dental pads, lips, or tongue. Filterability : Passes Gradocol membrane of about 400 millimicron average pore diameter. Other properties: Viable after at least 11 days at room temperature, 21 days at refrigerator temperature, 6 weeks frozen and dried in horse-serum saline. Literature: Mason and Neitz, Onder- stepoort Jour. Vet. Sci. and Anim. In- dust., 15, 1940, 159-173. Genus IV. Hostis gen. nov. Viruses of the Foot -and -]Mouth Disease Group, inducing diseases mainly charac- terized by vesicular lesions. Generic name from Latin hostis, enemy or stranger. The type species is Hostis pecoris spec. nov. Key to the species of genus Hostis. I. Infecting cattle and other animals with cloven hoofs; horse immune or highly resistant . II. Infecting horse readily. 1. Hostis pecoris spec. nov. From Latin pecus, cattle. Common names : Foot-and-mouth dis- ease virus; Virus der Maul- und Klauen- seuche. Hosts : Cow, pig, sheep, goat, reindeer, bison. Experimentall}^, also guinea pig, rabbit, rat. Insusceptible species: Chick embryo (chorioallantois) ; horse (immune or very resistant). Induced disease : In cow, after incuba- tion period of 2 to 4 days or more, fever, vesicular lesions on tongue, lips, gums, hard palate and feet, soon rupturing; salivation, lameness, generally recovery. 1. Hostis pecoris. 2. Hostis equinus. Transmission : Spread rapid, source of infection often obscure ; saliva is infective before lesions become obvious. Thermal inactivation : At 70° C-, not at 60° C, in 15 minutes. Filterability: Passes Seitz, Berkefeld V and X, and Chamberland Lu filters. Strains : Three strains. A, O and C, are iuimunologically distinct from each other. Other properties: Particle calculated to be about 20 millimicrons in diameter by centrifugation data, 8 to 12 millimicrons in diameter by filtration; may be sep- arated from mixtures with the larger equine vesicular stomatitis virus by dif- ferential filtration. Viable after drying 1240 MANUAL OF DETERMINATIVE BACTERIOLOGY in vacuo, at least a week at — 4 to 0° C. Readily destroyed by 1 to 2 per cent sodium hydrate or above pH 11. Soon inactivated near pH 6.0, but moderately stable at pH 2.0 to 3.0 ; optimum condition for storage at pH 7.5 to 7.7 in absence of air; return from 3.0 to 7.5 inactivates, however. Literature : Elford and Galloway, Brit. Jour. Exp. Path., 18, 1937, 155-161 ; Galloway and Elford, ibid., 14, 1933, 400- 408; 16, 1935, 588-613; 17, 1936, 187-204; Galloway and Schlesinger, Jour. Hyg., 37, 1937, 463-470; Hare, Jour. Path, and Bact., 35, 1932, 291-293; Loeffler and Frosch, Cent. f. Bakt., I Abt., 23, 1898, 371-391 ; Matte and Sanz, Bull. Soc. Path. Exot., 14, 1921, 523-529; Olitsky and Boez, Jour. Exp. Med., 45, 1927, 673-683, 685-699, 815-831, 833-848; Pyl, Ztschr. f. physiol. Chemie, 226, 1934, 18-28 ; Pyl and Klenk, Cent. f. Bakt., I Abt., Orig., 128, 1933, 161-171 ; Schlesinger and Gallo- way, Jour. Hyg., 37, 1937, 445-462. 2. Hostis equinus spec. nov. From Latin equinus, pertaining to horses. Common names: Vesicular-stomatitis virus, equine vesicular stomatitis virus. Hosts: Horse, domestic cattle. Ex- perimentally, also guinea pig, swine, white mouse, rabbit (relatively resistant ) , chick embryo ; Macaca mulatta (Zimmer- mann), rhesus monkey; M. irus, cynomolgus monkey. Insusceptible species : Chicken (ex- cept embryo ) . Geographical distribution : United States (Indiana, Xew Jersey). Induced disease: In horse, resembles foot-and-mouth disease of cattle; red- dened patches on buccal mucosa, mod- erate fever, .salivation, followed by ap- pearance of vesicles, especially on tongue, filled with clear or yellowish fluid; vesi- cles often coalesce and soon rupture leaving an eroded surface which heals soon in the absence of complications. Experimentally, in chorioallantois of de- veloping chick embryo, primary lesions involve moderate ectodermal prolifera- tion, degeneration, necrosis; mesodermal inflammation ; slight endodermal pro- liferation. Serological relationships: Strains iso- lated in different localities give antisera capable of neutralizing heterologous iso- lates of virus, but homologous antisera neutralize in higher dilutions than do heterologous antisera. Immunological relationships : No cross immunity with respect to equine en- cephalomyelitis virus. Filterability : Passes Seitz Alter. Other properties: May be separated from mixtures with foot-and-mouth dis- ease virus by propagation on chorioallan- toic membrane of chick embryo, which will not support increase of the latter virus. Inactivated by 1:50,000 methyl- ene blue in 2 mm layer 13 cm from 300 candle-power lamp in 60 minutes but not in 20 minutes. Particle estimated on the basis of filtration data to he 70 to 100 millimicrons in diameter; 60 millimicrons in diameter by centrifugation. Not de- stroyed by acidifying to pH 3 and return- ing to pH 7.5 (difference from foot-and- mouth disease virus). Literature : Burnet and Galloway, Brit. Jour. Exp. Path., 15, 1934, 105-113; Cox and Olitsky, Proc. Soc. Exp. Biol, and Med., 30, 1933, 053-654; Cox etal., ihid., 30, 1933, 896-898; Elford and Gallo- way. Brit. Jour. Exp. Path., 18, 1937, 155-161; Galloway and Elford, ibid., 14, 1933, 400-408; 16, 1935, 588-613; Mohler, Jour. Am. Vet. Med. Assoc, 52, 1918, 410-422; Olitsky et al.. Jour. Exp. Med., .59, 1934, 159-171 ; Pyl, Ztschr. f. physiol. Chemie, 226, 1934, 18-28; Sabin and Olitsky, ibid., 66, 1937, 15-34, 35-57; 67, 1938, 201-228, 229-249; Syverton ct al.. Science, 78, 1933, 216-217. FAMILY BORRELIOTACEAE 1241 Genus V. Molitor gen. nov. Viruses of the Wart-Disease Group, inducing diseases mainly characterized by tissue proliferation without vesicle or pustule formation. Generic name from Latin molitor, contriver. The type species is Molitor verrucae spec. nov. Key to the species of genus Molitor. I. Affecting man. II. Affecting cow. III. Affecting dog. IV. Affecting chicken. V. Affecting rabbit. 1. Molitor verrucae spec. nov. From Latin verruca, wart. Common name : Common-wart virus. Hosts -.HOMINIDAE — Homo sapiens L.,man. Perhaps also BOVIDAE — Bos taurus L., cow. CAN I DAE — Canis familiaris L., dog. Induced disease : Experimentally in man, incubation period long, 4 weeks to 6 or more months ; initially acanthosis (overgrowth of prickle cell laj'er of epi- dermis) and flattening of the papillae; later, interpapiliary hypertrophy, in- flammation, and marked hyperkeratosis. Transmission : By contact ; in some cases, venereally. Experimentally, bj^ skin scarification. Filterabilit}^ : Passes Berkefeld N filter. Literature : Ciuffo, Giorn. ital. d. malattie veneree e d. pelle, 48, 1907, 12- 17; Kingery, Jour. Am. Med. Assoc, 76, 1921, 440-442 ; Payne, Brit. Jour. Dermat., 3, 1891, 185-188; Schultz, Deutsch. med. Wchnschr., 34, 1908, 423; Serra, Giorn. ital. d. malattie veneree e d. pelle, 65, 1924, 1808-1814; Ullmann, Acta oto- laryngologica, 5, 1923, 317-334 ; Wile and Kingery, Jour. Am. Med. Assoc, 73, 1919, 970-973. 2. Molitor hominis comb. nov. {Strongyloplasma hominis Lipschiitz, 1. Molitor verrucae. 2. Molitor hominis. 3. Molitor bovis. 4. Molitor buccalis. 5. Molitor lumoris. 6. Molitor gingivalis. 7. Molitor sylvilagi. 8. Molitor myxomae. Arch. Dermat. u. Syph., 107, 1911, 395.) From Latin homo, man. Common name : Molluscum contagiosum virus. Host: HOMINIDAE— Homo sapiens L., man. Geographical distribution : Perhaps es- sentially world-wide. Induced disease : In man, experimen- tallj^, prodromal period may be 14 to 50 days, lesions at first like pimples, becom- ing red, painful, swollen, developing into small tumors covered with stretched and shiny skin; lesions commonest on face, arms, buttocks, back, and sides, heal- ing spontaneously. Inclusions within epithelial cells, known as molluscum bodies, measure 9 to 24 microns in diame- ter when approximately spherical, 24 to 27 microns in width and 30 to 37 microns in length when elongated; they contain elementary bodies about 0.3 micron in diameter. The outer envelope of the molluscum bodj^ is of carbohydrate. Transmission: By contact. By fo mites. Filterability : Passes Chamberland Li and Berkefeld V filters. Literature : Goodpasture and ffing. Am. Jour. Path., 3, 1927, 385-394; Good- pasture and Woodruff, ibid., 7, 1931, 1-8; Juliusberg, Deutsch. med. Wchnschr., 1242 MANUAL OF DETERMINATIVE BACTERIOLOGY 31, 1905, 1598-1599; Lipschiitz, Arch. Dermat. u. Syph., 107, 1911, 387-396; in Kolle, Kraus and Uhlenhuth, Handbuch der Pathogenen Mikroorganismen, 8, 1930, 1031-1040; Van Rooyen, Jour. Path, and Pact., 46, 1938, 425-436; 49, 1939, 345-349; Wile and Kingery, Jour. Cutan. Dis., 37, 1919, 431-446. 3. Molitor bovis spec. nov. From Latin bos, cow. Common name : Cattle-wart virus. Host: BOV IDAS— Bos taurus L., do- mestic cattle. Geographical distribution: United States. Induced disease: In cattle, especially about head, neck, and shoulders in young animals, on udders in cows, affected skin thickened at first, then rough, nodular; warts sometimes become large and pendu- lous, adversely affecting growth of host ; they sometimes become cauliflower-like tumors several inches in diameter; spon- taneous regression is not infrequent. Hides from affected animals are reduced in value. Transmission : Pelieved to be through injuries to skin when the injured part comes in contact with warty animals or with rubbing posts, chutes, fences, build- ings, or other structures with which affected animals have come in contact pre- viously. Experimentally, by skin inocu- lations, especially in animals under 1 ,vear of age. Filterability : Passes Perkefeld N filter. Literature: Creech, Jour. Agr. Res., 39, 1929, 723-737; U. S. Dept. Agr., Leaf- let 75, 1931, 1-4. 4. Molitor buccalis spec. nov. From Latin bucca, cheek. Common name : Canine oral-papillo- rnatosis virus. Host : CAN IDA E — Can is fa m il iar is L., dog. Insusceptible species: Cat, rabbit, guinea pig, rat, mouse; Macaca mulatta (Zimmermann), rhesus monkey. Induced disease : In young dog, experi- mentally, about 1 month after inocula- tion of buccal membrane by scarification, pale, smooth elevations, becoming gradu- ally^ more conspicuous and roughened ; finally a mass of closely packed papillae is formed. Regression with subsequent immunity is frequent ; no scars are left on regression. Secondary warts often appear in other parts of the mouth 4 to 6 weeks after primarj^ warts have first been observed. Transmission : Experimentally by skin scarification. Serological relationships : Not inhibited by antiserum effective against common- wart virus of man. Thermal inactivation : At some tem- perature between 45 and 58° C in 1 hour. Filterability : Passes Berkefeld N filter. Other properties : Viable after freezing and drying, if stored dry in icebox, at least 63 days ; in storage in equal parts of glycerine and 0.9 per cent NaCl solution at least 64 days. Literature : DeMonbreun and Good- pasture, Am. Jour. Path., 8, 1932, 43-56; M'Fadyean and Hobday, Jour. Comp. Path, and Therap., 11, 1898, 341-344; Penberthy, ibid., 11, 1898, 363-365. 5. Molitor tumoris spec. nov. From Latin tumor, swelling. Common names : Fowl-sarcoma virus, Rous chicken-sarcoma virus. Hosts : P HAS I AN I DAE— Callus gal- lus (L.), chicken. Experimentally, also pheasant (serial transfer difficult) and duck (by cell transfer only but filtrates from duck infect injected chicken). Insusceptible species : Turkey, guinea fowl (both immune to filtrates but ca- pable of supporting tumor line if alterna- ted in a series with common fowl hosts) ; geese . Induced disease : In hen, originally found in an adult, pure-bred hen of Barred Plymouth Rock variety. Experi- mentally transmitted, a circumscribed nodule soon becomes evident at site of implantation ; later this becomes necrotic or cystic at its center ; as growth enlarges, host becomes emaciated, cold, somnolent, and finally dies; discrete metastases are often found in lungs, heart, and liver. Parent cell of sarcoma is claimed to be FAMILY BORRELIOTACEAE 1243 the normal histiocj^te, but virus in the affected fowl is not confined to the sar- coma, being widespread in the body in spleen, liver, muscle, brain, etc. In the chick embryo, serial passage is feasible on the egg membrane, in which focal lesions involve only ectodermal tissue. Transmission : By injection of affected fowl cells or filtrates. Certain trans- missible tar-induced sarcomas, not in- fecting by filtrates, nevertheless induce the formation of antibodies capable of neutralizing this virus. An inhibitor of the virus extracted from tumors appears to be a protein, inactivated at 65° C, but not at 55° C, in 30 minutes and destroyed by trj'psin in 3 to 5 hours at pH 8. Oleic acid also may act as an inhibitor. No spontaneous transmission in chickens kept together. Serological relationships : Particles sedi- mented by centrifugal force 20,000 to 30,000 times gravity are specifically ag- glutinated by sera of fowls bearing corresponding tumor. At least one anti- gen in tumors of hen and duck not in healthy birds ; this one fixes complement and gives cross reactions with Rous, Mill Hill 2, Fujinami, and RFD2 tumors. Virus injected into goats produces two antibodies but only one if previously heated; the antibody to the heat-stable constituent requires complement to neu- tralize virus ; the only antibody produced in ducks does not require complement to neutralize. Thermal inactivation : At or below 54° C in 20 minutes. Filterability : Passes Berkefeld V and no. 5 (medium) filters. Other properties: Particle size esti- mated as about 100 millimicrons (but some say 50 or even 15 millimicrons) in diameter by filtration through graded membranes, about 70 millimicrons (molecular weight 140,000,000) by ultra- cent rifugat ion. Contains 8.5 to 9.0 per cent nitrogen, 1.5 per cent phosphorus. Protein tests positive. Feulgen reaction for thymonucleic acid absent ; 10 to 15 per cent of the protein maj^ be nucleic acid, probably of ribose type. PeBtose present . Virus believed to be of globulin nature or attached to globulin particles (Lewis and Mendelsohn, Am. Jour. Hyg., 12, 1930, 686-689). Viable indefinitely in dried spleen as in dried sarcoma tissues. Strains : Several strains have been studied in addition to the original Rous sarcoma no. 1 strain; immunological rela- tionships have been shown between the original strain, the des Ligneris sarcoma strain, the Fujinami sarcoma strain, the fibrosarcoma MHl and endothelioma MH2 strains; other isolates also have shown serological interrelationships. Literature : Amies, Jour. Path, and Bact., 44, 1937, 141-166; Amies et al.. Am Jour. Cancer, 35, 1939, 72-79 ; Andrewes Jour. Path, and Bact., 34, 1931, 91-107 35, 1932, 407-413 ; 37, 1933, 17-25, 27-44 43, 1936, 23-33; Claude, Jour. Exp. Med. 66, 1937, 59-72; Science, 87, 1938, 467-468 90, 1939, 213-214; Am. Jour. Cancer, 37 1939, 59-63; Claude and Rothen, Jour Exp. Med., 71, 1940, 619-633; Dmochow- ski and Knox, Brit. Jour. Exp. Path., 20, 1939, 466-472; Elford and Andrewes, ibid., 16, 1935, 61-66; Gye and Purdy, Jour. Path, and Bact., 34, 1931, 116-117 (Abst.) ; Haddow, ibid., 37, 1933, 149-155 ; Helmer, Jour. E.xp. Med., 64, 1936, 333- 338; Keogh, Brit. Jour. Exp. Path., 19, 1938, 1-9; Ledingham and Gye, Lancet, 228, 1935 {1), 376-377; Lewis and Men- delsohn, Am. Jour. Hyg., 12, 1930, 686- 689; des Ligneris, Am. Jour. Cancer, 16, 1932, 307-321 ; Mcintosh, Jour. Path, and Bact., 41, 1935, 215-217; Mellanby, Jour. Path, and Bact., 46, 1938, 447-460; 47, 1938, 47-64; Mendelsohn et al., Am. Jour. Hyg., 14, 1931, 421-425; Purdy, Brit. Jour. Exp. Path., 13, 1932, 473-479; Rous, Jour. Exp. Med., 13, 1911, 397-411. 6. Molitor gingivalis spec. nov. From Latin gingiva, gum. Common name : Rabbit oral -papilloma- tosis virus. Hosts: LEPORIDAE—Oryctolagm cuniculus (L.), domestic rabbit. Ex- perimentally, also Lepus americanus 1244 MANUAL OF DETERMINATIVE BACTERIOLOGY Erxlebeii, snowshoe hare; L. californicits Gray, jack rabbit ; Sylvilagus sp., cotton- tail rabbit . Geographical distribution: United States. Induced disease : In rabbit, benign papillomas, having the form of small, discrete, graj'-white, sessile or peduncu- lated nodules, usually multiple, on lower surface of tongue or, less frequently, on gums or floor of mouth. Transmission : Perhaps by mother to suckling young, with a latent period be- fore onset of disease. Not highly con- tagious, if contagious at all, in old animals. Experimentally by puncture of tissues in the presence of virus. Immunological relationships : Specific immunitj' develops as a result of disease, but no cross immunity with respect to rabbit-papilloma virus, which differs also in failing to act on oral mucosa. Filterabilily : Passes Berkefeld V and N filters. Literature: Parsons and Kidd, Jour. Exp. Med., 77, 1943, 233-250. 7. Molitor sylvilagi spec. nov. From Xew Latin Sylvilagus, generic name of cottontail rabbit. Common names : Rabbit papilloma or papillomatosis virus, rabbit wart virus. Hosts: LEPORIDAE— Sylvilagus sp., cottontail rabbit. Experimentally, also LEPORIDAE — Oryctolagus nmiadits (L.), domestic rabbit. Geographical distribution : United States. Induced disease : In cottontail rabbit, at first minute elevations along lines of scarification; later solid masses of wrin- kled keratinized tissue, 3 to 4 millimeters in thickness; eventually cornified warts, striated perpendicularly at top, fleshy at base, 1 to 1.5 cm in height; regression rare ; natural papillomas become malig- nant occasionally. In domestic rabbit, experimentally, blood antibody remains low but virus is always masked, prevent- ing serial passage ; discrete lesions on skin permit quantitative tests ; tarring causes localization of virus from blood stream; papillomas give rise to malignant acan- thomatous tumors by graded continuous alteration; metastasis frequent; trans- plantation to new hosts successful in series ; antibody specific for the virus is formed continuously in the transplanted growths although virus is not directly demonstrable by subinoculation from them; malignant growths appear more promptly and frequently where epidermis has been tarred long; virus appears specific for epithelium of skin; growths disappear if treated with X-raj'S, 3600 r at one time or fractionally; 60 per cent are cured with 3000 r, but 2000 r ineffec- tive. Transmission : Experimentally, by scarification of skin. Abnormal suscepti- bility to infection is noted in rabbit skin treated with 0.3 per cent methylcholan- threne in benzene or equal parts of tur- pentine and acetone. Serological relationships : Specific neu- tralization, reversible on dilution. Com- plement fixation specific, with virus particle as antigen ; no cross reaction with antisera for vaccinia, herpes, fibroma, or myxoma viruses. Precipitates occur in properly balanced mixtures of virus and specific antiserum; virus and antibody in both free and neutralized states are present in both soluble and insoluble phases of these suspensions. Immunological relationships : Intra- peritoneal injections immunize specifi- cally. Rabbits immunized to fibroma and myxoma viruses are susceptible to rabbit papilloma virus. Thermal inactivation : At 70° C, not at 65 to 67°C, in 30 minutes; in 0.9 per cent sodium chloride solution at 65 to 66° C, time not stated. Filterability : Passes Berkefeld V, N, and W filters ; particle size calculated as 23 to 35 millimicrons by filtration as com- pared with 32 to 50 millimicrons by cen- trifugation and 44.0 millimicrons by measurement of electron micrographs, which show the particle to be approxi- mately spherical in shape. FAMILY BOKRELIOTACEAE 1245 Other properties: Infectious particle has sedimentation constant S20 = ca. 250 X 10~^' cm per sec. per dyne; usually there is a secondary boundary at about 375 X 10~i^. Isoelectric point between pH 4.8 and 5.1. Maximum absorption at about 2750 A. Contains thj-mus nucleic acid about 6.8 to 8.7 per cent ; maximum absorption of nucleic acid at about 2630 A. Literature: Beard et al., Jour. Inf. Dis., 65, 1939, 43-52; 69, 1941, 173-192; Bryan and Beard, ibid., 65, 1939, 306-321 ; Friedewald, Jour. Exp. Med., 75, 1942, 197-220; Hoyle, Jour. Path, and Bact., 50, 1940, 169-170; Kidd, Jour. Exp. Med., 68, 1938, 703-724, 725-759; 70, 1939, 583- 604; 71, 1940, 469-494; 74, 1941, 321-344; 75, 1942, 7-20; Kidd and Rous, ibid., 68, 1938, 529-562; 71, 1940, 813-838, Kidd et al., ibid., 64, 1936, 63-77, 79-96; Rous and Beard, ibid., 60, 1934, 701-722; 62, 1935, 523-548; Rous and Kidd, ibid., 67, 1938, 399-428; 71, 1940, 787-812; Rous et al., ibid., 64, 1936, 385-400, 401-424; Schles- inger and Andrewes, Jour. Hyg., 37, 1937, 521-526; Sharp et al., Proc. Soc. Exp. Biol, and Med., 50, 1942, 205-207; Shope, Jour. Exp.IMed., 58, 1933, 607-624 ; 65, 1937, 219-231; Syverton et al., ibid., 73, 1941, 243-248; Taylor et al., Jour. Inf. Dis., 71, 1942, 110-114. 8. Molitor myxomae (Aragao) comb, nov. (Chlamidozoon myxomae Aragao, Brazil-Med., 25, 1911, 471; name later abandoned by its original author in favor of Strongyloplasma myxomae Aragao, Mem. Inst. Oswaldo Cruz. 20, 1927, 231 and 243. The name SanarelUa cnnicidi Lipschiitz, Wien. klin. Wochenschr., 40, 1927, 1103, was based on the supposed causative organism, defined as varying in size between the size of chlamydozoa and of large cocci ; it is not clear whether the structures observed and named were virus particles or not.) From New Latin viyxoma, a kind of soft tumor, from nature of induced lesion. Common names : M3^xoma virus, virus myxomatosuvi . Hosts : LEPORIDAE—Oryclolagus cuniculiis (L.), domestic rabbit. Ex- perimentally, also Sylvilagus sp., cotton- tail rabbit ; jack rabbit (once in many trials); Lepus brasiliensis (resistant and rarely infected). Also chick embiyo and duck embryo. Insusceptible species : Lepus califor- nicus Gray, black-tailed jack rabbit; L. americanus Erxleben, varying hare ; Syl- vilagus transitional is Bangs, cottontail; horse, sheep, goat, cattle, dog (but one reported infected), guinea pig, rat, mouse, fowl, pigeon, duck, cat, hamster, monkey; man (but some conjunctival pain and swelling). Geographical distribution : South Amer- ica (Brazil, Uruguay, Argentina), United States (California). Induced disease : In domestic rabbit, a disease {myxomatosis cuniculi) almost al- waj's fatal at ordinary room temperatures but not at 36 to 42° C, lesions fewer and regressing after 6 to 8 days at these higher temperatures in most animals. At ordi- nary temperatures, nodules (edematous tumors) in skin near eyes, nose, mouth, ears, and genitalia; edema of eyelids; conjunctivitis with purulent discharge if skin around eyes is involved. Later marked dj'spnea, stertorous breathing, cyanosis, asphy.xia. Animals usually die within 1 to 2 weeks of infection. Virus enters bloodstream and invades nervous system at random through walls of blood vessels. Discharges from nose, eyes, and the serous exudates from affected tissues are infectious; urine and feces are not. There are cytoplasmic inclusions in af- fected epidermal cells. In chick em- bryo, experimentally, intense inflamma- tion, eventual impairment of circulation and necrosis locally ; growth best if em- brj^o is grown at 33 to 35° C and chilled to 25° C for 12 to 18 hours before or after inoculation, lesions being linear and asso- ciated with capillaries in ectoderm; virus infects and is recoverable from embryo and depresses hatch. Transmission : By contact with dis- eased rabbits or cages recently occupied 1246 MANUAL OF DETERMINATIVE BACTERIOLOGY by them. Through air for a few inches. Rarely by feeding. Experimentally, by rubbing conj unctiva with a bit of infected tissue or with a platinum loop contam- inated from diseased conjunctiva; has been recovered from flies. By injection. By flea, Clenopsyllafelis iPULICIDAE), rarely. Serological relationships : An attack of the disease induces the formation of neu- tralizing antibodies. Cross neutraliza- tion by antisera to myxoma and fibroma strains. Complement is fixed with myx- oma virus as test antigen in the presence of antisera to myxoma or fibroma strains. Serum of rabbit inoculated with a soluble antigen, a heat-labile protein with iso- electric point near pH 4.5, agglutinates myxoma elementary bodies. A second soluble antigen, also heat labile, appears distinct, inhibiting its own antibody even after inactivation of its precipitating power by exposure at 56° C. Immunological relationships : Myxoma- recovered domestic rabbits become immune to reinfection; fibroma-strain-re- covered animals, although partially im- munized, still support myxoma-strain virus introduced into the testicle. Heat- inactivated virus (60° C for 30 minutes) tends to immunize if given intradermally ; there is then an allergic local response, less severe generalized disease, delayed death or recovery. If fibroma virus pre- cedes myxoma virus by 48 to 96 hours, there is marked protection. Thermal inactivation: At 55° C in 10 minutes ; at 50° C in 1 hour. A substance thermostabile for 30 minutes at 60 to 75° C, but not at 90° C, is itself unable to produce myxomatous changes after the heat treatment but may do so in com- bination with fibroma virus, and trans- missible myxoma virus is then reconsti- tuted. Although it is supposed by some that this indicates the transformation of fibroma-strain virus into myxoma-strain virus, the possibility that heat-modified myxoma-strain virus is reactivated has not been eliminated. Filterability : Passes Berkefeld V and N filters ; not Chamberland L5 or L7 filters. Other properties : Inactivated above pH 12.0 and below pH 4.0. Withstands drying. Viable at least 3 months at 8 to 10° C. Literature : Aragao, Brazil-med., B5, 1911, 471; Mem. Inst. Oswaldo Cruz,^0, 1927, 225-235; Berry and Dedrick, Jour. Bact., 31, 1936, 50-51 (Abst.) ; Berry and Lichty, ibid., 31, 1936, 49-50 (Abst.); Berry et al.. Second International Cong- ress for Microbiology, Report of Proceed- ings, London, 1936, 96 (Abst.) ; Fisk and Kessel, Proc. Soc. Exp. Biol, and Med. 29, 1931, 9-11 ; Gardner and Hyde, Jour Inf. Dis., 71, 1942, 47-49; Hobbs, Am Jour. Hyg., 8, 1928, 800-839; Science, 73 1931, 94-95; Hoffstadt and Omundson Jour. Inf. Dis., 68, 1941, 207-212; Hoff stadt and Pilcher, Jour. Bact., 35, 1938 353-367; 39, 1940, 40-41 ; Jour. Inf. Dis. 64, 1939, 208-216; 65, 1939, 103-112 Hoffstadt et al., ibid., 68, 1941, 213-219 K. E. Hyde, Am. Jour. Hyg., 23, 1936 278-297; R. R. Hyde, tbid., 30 (B), 1939 37-46, 47-55; Hyde and Gardner, ibid. 17, 1933, 446-465; 30, (B), 1939, 57-63 Kessel et al., Proc. Soc. Exp. Biol, and Med., 28, 1931, 413-414; Lipschutz Wien. klin. Wchschr., .^0, 1927, 1101-1103 Lush, Austral. Jour. Exp. Biol, and Med Sci., 15, 1937, 131-139; 17, 1939, 85-88 Martin, Austral. Counc. Sci. and Indust Res., Bull. 96, 1936, 28 pages; Moses Mem. Inst. Oswaldo Cruz, 3, 1911, 46-53 Parker and Thompson, Jour. Exp. Med. 75, 1942, 567-573; Plotz, Compt. rend Soc. Biol., Paris, 109, 1932, 1327-1329 Rivers, Jour. Exp. Med., 51, 1930, 965- 976; Rivers and Ward, ibid., 66, 1937 1-14; Rivers et al., ibid., 69, 1939, 31-48 Sanarelli, Cent. f. Bakt., I Abt., 23 1898, 865-873; Shaffer, Am. Jour. Hyg. 34 (B), 1941, 102-120; Shope, Jour. Exp Med., 56, 1932, 803-822; Smadel et al. ibid., 72, 1940, 129-138; Splendore, Cent f. Bakt., I Abt., Orig., 48, 1909, 300-301 Stewart, Am. Jour. Cancer, 15 Suppl., FAMILY BORRELIOTACEAE i247 1931, 2013-2028; Swan, Austral. Jour. Exp. Biol, and Med. Sci., 19, 1941, 113- 115. Strains and substrains : A strain from cottontail rabbits (Sylvilagus sp.), differ- ing from typical myxoma virus, has been studied extensively under the name fibroma virus. This strain in turn is rec- ognized as consisting of variants and has been investigated as typical (OA) and inflammatory (lA) substrains, antigeni- cally alike but the latter tending to generalize in domestic rabbits. Fibroma virus is not lethal in domestic rabbits as the type strain almost always is; it appears to lack some antigenic constitu- ents, inducing the formation of agglu- tinins that give cross reactions with the type but of neutralizing and complement - fixing antibodies that do not. The fi- broma strain does not generally appear in the blood stream, as myxoma virus does, and is not contagious, at least it does not spread spontaneously among domestic rabbits as the myxoma strain does; the manner of its spread in wild rabbits in nature is not known. Its particle size has been calculated as 126 to 141 millimicrons by centrifugation, 125 to 175 millimicrons by filtration. (Ahlstrom, Jour. Path, and Bact., 4(>, 1938, 461-472 ; Andrewes, Jour . Exp. Med., 63, 1936, 157-172 ; Hoffstadt and Pilcher, Jour. Inf. Dis., 68, 1941, 67-72; Hurst, Brit. Jour. Exp. Path., 18, 1937, 1-30; Austral. Jour. Exp. Biol, and Med. Sci., 16, 1938, 53-64, 205-208; Hyde, Am. Jour. Hyg., 24, 1936, 217-226; Ledingham, Brit. Jour. Exp. Path., 18, 1937,436-449; van Rooyen, ibid., 19, 1938, 156-163; van Rooyen and Rhodes, Cent. f. Bakt., I Abt.,0rig.,U:2, 1938, 149-153 ;Schlesinger and Andrewes, Jour. Hyg., 37, 1937, 521-526; Shope, Jour. Exp. Med., 66, 1932, 793-822; 63, 1936, 33-41, 43-57, 173-178. 1248 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY III. ERRONACEAE FAM. NOV. Viruses of the Encephalitis Group, inducing diseases mainly characterized by ef- fects on nerve tissues. Key to the genera of family Erronaceae. I. Viruses of the Typical Encephalitis Group. Genus I. Erro, p. 1248. II. Viruses of the Poliomyelitis Group. Genus II. Legio, p. 1257. III. Viruses of the Rabies Group. Genus III. Formido, p. 1263. Genus I. Erro gen. nov. Viruses of the Typical Encephalitis Group, inducing diseases mainly characterized by injuries to cells of the brain. Vectors of some known to be ticks ; dipterous insects may also transmit. Generic name from Latin erro, a vagrant. The type species is Erro scoticus spec. nov. Key to the species of genus Erro. I. Affecting sheep principally, but also man. 1. Erro scoticus. II. Affecting man principally. 2. Erro silvesiris. 3. Erro incognitus. 4. Erro japonicxis. 5. Erro nili. 6. Erro scelestus, III. Affecting horse principally, but also man. 7. Erro equinus. IV. Affecting horse, cow, sheep. 8. Erro bornensis. 1. Erro scoticus spec. no;.'. FromLatin lion, champingof jaws; prostration, coma, Scoticus, Scottish. death. In man, encephalitis with Common name: Louping-ill virus. prompt and complete recovery accom- Hosts: BOVIDAE—Ovis aries L., panied by formation of specific neutraliz- sheep. HOM I N I DAE— Homo sapiens ing antibodies. In mouse, experimen- L., man. Experimentally, also mouse, tally, diffuse encephalomyelitis with mild rat (subclinical infection), chick embryo meningeal involvement; following intra- (discrete primary lesions on chorioallan- cerebral inoculation, fine rhythmical toic membrane), Macacus rhesus, horse, tremor involving neck, nose, and ears, cow, pig. unsteadiness, muscle spasms, respiratory Insusceptible species : Guinea pig, distress, sometimes clonic and rarely rabbit. tonic convulsions; hind limb paralysis, Geographical distribution : Scotland, dribbling of urine, cessation of spontane- northern England. ous limb movements, death; in mouse Induced disease: In sheep, encephalitis inoculated intraperitoneally, virus usu- characterized by dullness followed by ally enters central nervous system by incoordination of movement, frequently way of the olfactory mucosa and olfactory with tremors chiefly of the head; saliva- bulb, occasionally by trauma at points of FAMILY ERRONACEAE 1249 damage ; in mouse inoculated iutranasally , virus enters blood and reaches the olfac- tory bulb where it multiplies to a high concentration before infecting the re- mainder of the brain and the rest of the nervous system ; tends to disappear from the blood after sickness begins but per- sists in the brain until death from encephalitis. In chick embryo, after inoculation of chorioallantoic membrane, edema and opacity spreading from site of inoculation on membrane of 10-day embryo ; in 12-day eggs, discrete primary lesions, sometimes with secondary lesions surrounding them on the inoculated mem- brane ; embryo dies in about 6 days, after showing jaundice, edema, mottling of the liver with necrosis ; virus regularly in blood. In monkey, Macacus rhesus, progressive cerebellar ataxia ; encephalo- myelitis with involvement and massive destruction of Purkinje cells in the cere- bellum. Transmission : By ticks, Rhipicephalus appendiculatus and Ixodes ricinus (IXO- DIDAE). In Rhipicephalus appendicu- latus, the larva or nymph becomes in- fected ; only a few individuals retain virus until the adult stage ; virus does not pass through the egg. Non-viruliferous ticks do not acquire virus by feeding with infective ticks on immune animals. Ex- perimentally, by intracerebral or intra- peritoneal injection in mouse ; by intra- nasal instillation in rat, mouse, and monkey. Serological relationships : Complement fixation and neutralization tests show cross reactions with Russian spring- summer encephalitis virus, but immune serum against louping-ill virus is only partially effective in neutralizing the spring-summer encephalitis virus. Immunological relationships: Mice are protected against louping-ill virus by vaccination with non-virulent spring- summer encephalitis virus but protection is less effective than for the homologous virus. No cross immunity with respect to Rift Valley fever virus or poliomyelitis virus in Macacus rhesus, but immunity with respect to reinfection by louping-ill virus has been demonstrated. Thermal inactivation : At 58° C in 10 minutes. Filtefability : Passes Berkefeld V, N, and W filters. Other properties : Viable in broth fil- trates after storage at 4° C and pH 7.6 to 8.5 for 70 days. Particle diameter, cal- culated from ultrafiltration data, 15 to 20 millimicrons. Literature : Alexander and Neitz, Vet. Jour., 89, 1933, 320-323; Onderstepoort Jour. Vet. Sci. and Anim. Industr., 5 1935, 15-33; Alston and Gibson, Brit Jour. Exp. Path., 1£, 1931, 82-88; Burnet Jour. Path, and Bact., 4.2, 1936, 213-225 Brit. Jour. Exp. Path., 17, 1936, 294-301 Burnet and Lush, Austral. Jour. Exp Biol, and Med. Sci., 16, 1938, 233-240 Casals and Webster, Science, 97, 1943 246-248; Jour. Exp. Med., 79, 1944 45-63; Elford and Galloway, Jour. Path and Bact., 37, 1933, 381-392; Findlay Brit. Jour. Exp. Path., 13, 1932, 230-236 Fite and Webster, Proc. Soc. Exp. Biol, and Med., 31, 1934, 695-696; Galloway and Perdrau, Jour. Hyg., 35, 1935, 339- 346; Hurst, Jour. Comp. Path, and Therap., U, 1931, 231-245; M'Fadyean, Jour. Comp. Path, and Therap., 7, 1894, 207-219; 13, 1900, 145-154; Pool et al., ibid., 43, 1930, 253-290; Rivers and Schwentker, Jour. Exp. Med., 59, 1934, 669-685; Schwentker et al., ihid., 57, 1933, 955-965. 2. Erro silvestris spec. nov. From Latin silvestris, of the forest, in reference to incidence of the induced disease almost exclusivel}' in those who enter forest lands. Common names : Spring-summer en- cephalitis virus, forest spring encephalitis virus. Hosts: Man; probably cattle, horse; Euiamias asiaticus orientalis, Evotomys rufocanus arsenjevi. Experimentally, also white mouse, Macacus rhesus, birds, goat, sheep, Microius michnoi pelliceus Thom., Cricetuhis furunculus. 1250 MANUAL OF DETERMINATIVE 6ACTEM0L0GY Geographical distribution : Union of Soviet Socialist Republics. Induced disease: In man, acute non- suppurative encephalitis, abrupt onset, steep rise of temperature to 38 to 40° C, severe headache, giddiness, and vomit- ing; pareses and paralyses of upper or lower limbs or muscles of neck and back ; residual atrophic paralyses common; mortality among cases, 30 per cent ; 80 per cent of all cases occur in May and June. Transmission : By tick, Ixodes persul- catus (IXODIDAE) ; the virus seems to hibernate in this species and has proved capable of passing through eggs to prog- eny. Experimentally, also by ticks Dermacentor silvarum and Haemaphysa- lisconcinna (IXODIDAE). Serological relationships : Virus-neu- tralizing antibodies, found without other evidence of disease in some men and in many cattle and horses, believed to indi- cate susceptibility of these hosts to latent infections. No cross neutralization with St. Louis encephalitis virus. Japanese summer encephalitis virus is in part antigenically related, but some antigenic constituents of this virus are missing in spring-summer encephalitis virus and vice versa. Immunological relationships: Formol- ized virus immunizes specifically. Filterability : Passes Berkefeld and Chamberland filter candles. Literature : Smorodintseff, Arch, f . gesamt. Virusforsch., 1, 1940, 468-480; Soloviev, Acta Med. U. R. S. S., 1, 1938, 484-492 (Biol. Abst., 17, 1943, 1726, no. 18777). 3. Erro incognitus spec. nov. From Latin incognitus, unknown, in reference to mystery surrounding the nature and relationships of this virifs, as evidenced by common name. Common name : Australian X-disease virus. Hosts : HOMINIDAE — Homo sapiens L., man. Experimentally, also sheep, horse, cow, rhesus monkey. Geographical distribution : Australia. Induced disease : In man, polioenceph- alitis, especially in children, occurring in late summer; mortality high; charac- terized by headache, body pains, drowsi- ness, weakness, then vomiting, fever, convulsions; paralysis of limbs, eye- muscles, or face rare; recovery rapid in non-fatal cases. Literature: Kneebone, Austral. Jour. Exp. Biol, and Med. Sci., 3, 1926, 119- 127; Perdrau, Jour. Path, and Bact., 4^, 1936, 59-65. 4. Erro japonicus spec. nov. From New Latin Japonia, Japan. Common name : Japanese B encepha- litis virus. Hosts : HOMIN IDAE—Homo sapiens L., man. Experimentally, also young sheep, mouse, and Macacus rhesus. Geographical distribution: Japan, Union of Soviet Socialist Republics. Induced disease : In man, loss of appe- tite, drowsiness, nausea, then rapid rise of temperature, pains in joints and chest ; restlessness followed by apathy, coma; death, usually before end of second week, or recovery, sometimes with persistence of evidences of damage done to the nerv- ous sj^stem by the disease. Serological relationships : Specific anti- serum does not neutralize St. Louis en- cephalitis virus or louping-ill virus. Rus- sian autumn-encephalitis virus induces the formation of antisera neutralizing Japanese B encephalitis virus. Russian spring-summer encephalitis virus con- tains some, but not all, antigens in com- mon with this virus. Australian X-dis- ease virus is distinct in neutralization tests. Immunological relationships : Specific immunity as a result of earlier infection in mice ; no cross protection with respect to St. Louis encephalitis virus. Vac- cination with Japanese B encephalitis virus does not enhance resistance to West Nile encephalitis virus but only to the homologous virus. FAMILY EREONACEAE 1251 Thermal inactivation : At 56° C in 30 minutes. Filterability : Passes Berkefeld N, W, Chamberland L2, Lg, and Seitz EK filters, with ease. Literature: Kudo et al., Jour. Im- munol., 32, 1937, 129-135; Smorodintseff et al.. Arch. f. gesamt. Virusforsch., 1, 1940, 550-559 ; Webster, Jour. Exp. Med., 67, 1938, 609-618. 5. Erro nili spec. nov. From Latin Nilus, god of the Nile. Common name : West Nile encephalitis virus. Hosts : HOMINIDAE— Homo sapiens L., man (perhaps without inducing any definite disease). Experimentally, also rhesus monkey, mouse. Geograpliical distribution : Africa (Uganda). Induced disease : In man, no details are known ; virus was originally isolated from blood of a woman native of Uganda ; at the time the temperature of the patient was 100.6° F but she denied illness ; moreover, two laboratory workers developed neu- tralizing antibodies without recognizable clinical disease. In mouse, experimen- tally, after intracerebral inoculation, in- cubation period to 4 or 5 days, then hyperactivity and roughening of coat ; later, weakness, hunched attitude, some- times paralysis of hind quarters ; usually coma before death. In rhesus monkey, experimentally, after intracerebral or intranasal inoculation, fever and en- cephalitis. Serological relationships : No cross reac- tions in complement fixation tests between this and equine encephalitis virus, Japanese B encephalitis virus, St. Louis encephalitis virus, or lymphocytic chori- omeningitis virus. Neutralization tests show some common antigens in West Nile encephalitis virus, Japanese B encepha- litis virus and St. Louis encephalitis virus ; antiserum to West Nile virus does not neutralize either of the others but antisera against St. Louis virus may neutralize West Nile virus and antisera against Japanese B virus have some effec- tiveness in neutralizing both West Nile virus and St. Louis virus. Immunological relationships : Vaccina- tion with this virus does not enhance resistance to Japanese B or St. Louis encephalitis viruses but only resistance to the homologous virus. Thermal inactivation : At 55° C, not at 50° C, in 30 minutes. Filterability: Passes Berkefeld V, N, and W filter candles readily; also passes Seitz EK asbestos pads and collodion membranes 79, not 62, millimicrons in average pore diameter. Other properties : Infective particle 21 to 31 millimicrons in diameter, as calcu- lated from filtration experiments. Vi- able at least 2 weeks at 2 to 4° C. Viable after drying from the frozen state. Literature: Havens et al.. Jour. Exp. Med., 77, 1943, 139-153; Smithburn, Jour. Immunol., U, 1942, 25-31 ; Smithburn et al.. Am. Jour. Trop. Med., 20, 1940, 471- 492. 6. Erro scelestus spec. nov. From Latin scelestus, infamous. Common name: St. Louis encephalitis virus. Hosts : HOMINIDAE — Homo sapiens L., man. A great number of mammals and birds in endemic areas may have antisera that neutralize the virus, indi- cating that they are probably natural hosts; among these are : ANATIDAE— Anas plalyrhyncha L., Mallard and Pekin ducks; Anser anser (L.), domestic goose. BOVIDAE — Bos taurus L., cow; Capra hircus L., goat; Ovis aries L., sheep. CANIDAE — Canis familiaris L., dog. COL UMBIDAE—Columba livia, domes- tic pigeon ; Zenaidura macroura, western mourning dove. EQUIDAE — Equus ca- ballvs L., horse. FALCONIDAE— Falco sparverius L., sparrow hawk. LE- PORIDAE — Lcpiis californicus Gray, jack rabbit ; Sylvilagus nuiialli, cotton- tail rabbit. MELEAGRIDAE—Mele- agris gallopavoh., turkey. MURIDAE — Rattus norvegicus (Berkenhout), brown 1252 MANUAL OF DETERMINATIVE BACTERIOLOGY rat. MUSCICAPIDAE—Tiirdus mi- gratorius L.,Tohm. PHASIANIDAE— Gallus gallus (L.), chicken; Lophortyx calif ornica, California quail. PICIDAE — Asyndesmus lewis, Lewis woodpecker; Colaptes cafer (Gm.), red-shafted flicker. STRIGIDAE— Bubovirginianus (Gm.), great horned owl. S UIDAE—Svs scrofa L., pig. Experimentally, white mouse (some substrains of the Swiss white mouse are genetically more readily in- fected than others); Macacus rhesvs; pigeon (inapparent infection) ; chick embryo and to a limited extent the young hatched chick. Insusceptible species : Laboratory rab- bit, Cebus monkey, guinea pig, rat. Geographical distribution : L'nited States. Induced disease : In man, during sum- mer and fall, about 9 to 21 days after ex- posure, headache, high fever, rigidity of neck, tremors; encephalitis, usually with fever; some patients become drowsy, others sleepless or delirious; usual se- quelae headaches, irritability, some loss of memory, and drowsiness ; neutralizing antibodies maintained in vivo at least 2^ years after occurrence of disease. Experimentally, in susceptible strains of white mouse inoculated by intracerebral injection, after 3 to 4 days, coarse tremors, convulsions, prostration, death; perivas- cular accumulations of mononuclear leuco- cytes throughout brain, stem, cord, and pia, with destruction of pyramidal cells in the lobus piriformis and cornu Am- monis ; subcutaneous and intraperitoneal injections immunize against subsequent infection by intracerebral inoculation, virus reaching only blood and spleen in the process of immunization unless an excessive dose is given; some substrains of the White Swiss mouse are relatively resistant to infection, requiring inocula- tion with about 1000 times the minimal infective dose for highly susceptible strains and when infected proving rela- tively poor sources of virus for subinocu- lation; highly susceptible substrains of the White Swiss mouse lack a single major, dominant, genetic factor that is present in resistant substrains. Transmission : By mosquito, Cnlex tar- salis Coquillett {CULICIDAE), prob- ably extensively; this insect has been collected in nature carrying the virus. Experimentally, by larvae of American dog tick, Dcrmacentor variabilis (Say) ilXODIDAE); by mosquito, Culex pipiens Linn., var.pallans Coq. (CULI- CIDAE). To mice, by feeding on in- fected tissues. Serological relationships : Human anti- sera may neutralize virus after clinical and subclinical attacks. Immunological relationships : Specific intracerebral immunity after vaccination by subcutaneous or intraperitoneal injec- tion in mice appears early (about 1 week after vaccination) and disappears before humoral antibody titer reaches its max- imum. Thermal inactivation : At 56° C in 30 minutes. Filterability : Passes Berkefeld V and N filter candles and collodion membranes 66 millimicrons in average pore diameter. Other properties : Storage in human brain tissue in glycerine inactivates this virus in about 32 days. Diameter of infective particle calculated from filtra- tion data as about 20 to 33 millimicrons. In storage, rabbit and sheep sera act to some extent as preservatives. At 4° C, after drying in vacuo while frozen, viable in apparently undiminished titer for at least 17 months. Literature : Bang and Reeves, Jour. Inf. Dis., 70, 1942, 273-274 ; Bauer et al., Proc. Soc. Exp. Biol, and Med., 31, 1934, 696-699; Blattner and Cooke, Jour. Inf. Dis., 70, 1942, 226-230; Blattner and Heys, Proc. Soc. Exp. Biol, and Med., 48, 1941, 707-710; Cook, .Jour. Inf. Dis., 63, 1938, 206-216; Cook and Hudson, ibid., 61, 1937, 289-292; Elford and Perdrau, Jour. Path, and Bact., 40, 1935, 143-146; Hammon and Howitt, Am. Jour. Hyg., 35, 1942, 163-185; Hammon et al.. Science, 94, 1941, 305-307, 328-330; Jour. Inf. Dis., 70, 1942, 263-266, 267-272, 278- FAMILY ERRONACEAE 1253 283; Harford and Bronfenbrenner, Jour. Inf. Dis., 70, 1942, 62-68; Harrison and Moore, Am. Jour. Path., 13, 1937, 361- 375; Hodes, Jour. Exp. Med., 69, 1939, 533-543; Hodes and Webster, ihid., 68, 1938, 263-271 ; Lennette and Smith, Jour. Inf. Dis., 65, 1935, 252-254 ; Mitamura et al., Trans. Soc. Path. Jap., 27, 1937, 573- 580; Muckenfuss et al., U. S. Pub. Health Service, Public Health Kept., 48, 1933, 1341-1343; O'Leary et al., Jour. Exp. Med., 75, 1942, 233-246; Reeves et al., Proc. Soc. Exp. Biol, and Med., 50, 1942, 125-128; Sulkin et al.. Jour. Inf. Dis., 67, 1940, 252-257; Webster, Jour. Exp. Med., 65, 1937, 261-286; 6S, 1938, 111-124; Web- ster and Clow, ibid., 63, 1936, 433-448, 827-845; Webster and Fite, ibid., 61, 1935, 103-114, 411-422; Webster and Johnson, ibid., 7 It, 1941, 489-494 ; Webster et al., 61, 1935, 479-487; 6,2, 1935, 827-847. 7. Erro equinus spec. nor. From Latin equinus, pertaining to horses. Common name: Equine encephalitis virus. Hosts: EQUIDAE—Equus caballus L., horse ; Fi hybrid of the horse and E. asinus L.,mule. HOMINIDAE—Homo sapiens L., man. COLUMBIDAE — Columba livia, domestic pigeon. PHA- SIA N IDAE — ring-necked pheasant. TETRAONIDAE—Tympanuchus cup- ido L., var. americanus (Reichenbach), prairie chicken. Many additional spe- cies have been found to show neutralizing antisera at times and these are presum- ably natural hosts of the virus upon occa- sion; among them are: ANATIDAE — Anas platyrhyncha L., Mallard and Pekin ducks; Anser anser (L.), domestic goose. BOVIDAE — Bos laurus L., cow ; Capra hircus L., goat ; Ovis aries L., sheep. CAN I DAE — Canis familiaris L., dog. CHARADRIIDAE—Oxyechusvociferus L., killdeer. CRICETIDAE—Microtus montanus (Peale), field mouse; Peromys- cus vianiculatus (Wagner), white-footed mouse.' FALCONIDAE — Falco spar- rerius L., sparrow hawk. MELEAGRI- DAE — Meleagris gallopavo L., turkey. MURIDAE—Rattus rattus L., black rat. MUSCICAPIDAE—Turdus mi- gratorius L., robin. MUSTELIDAE— Mustela frenata Lichtenstein, weasel. PHASIANIDAE—Gallus gallus (L.), chicken; Lophortyx calif ornica, Cali- fornia quail ; Phasianus colchicus L., ring- necked pheasant. PIC I DAE — Colaptes cafer (Gm.), red-shafted flicker. STRI- GIDAE — Bubo virginianus (Gm.), great horned owl. SUIDAE — Sus scrofa L., pig. Experimentally, also chick embryo, goose embryo, pheasant embryo, robin embryo, pigeon embryo, turkey embryo, sparrow embryo, duck embrj'o, and gui- nea-fowl embryo; white mouse, guinea pig, rabbit, pigeon, white rat, calf, sheep, monkey, goat, dog, hen, turkey; Zono- trichia leucophrys gambeli, Gambel spar- row; Passer domesiicus L., English sparrow; Lophortyx calif ornica, quail; Junco oreganus, junco; Toxosioma lecon- iei lecontei, thrasher ; Citellus richardsonii (Sabine), gopher or Richardson's ground squirrel ; Sigmodon kispidus Say and Ord, cotton rat ; Dipodomys heermanni Le ■ Conte, kangaroo rat; Reithrodoniomys megalatus, wild mouse; Microtus mon- tanus, M. californicus and M. mordax, wild mice ; Peromyscns maniculatus (Wag- ner), white-footed mouse; Neotoma fus- cipes Baird, wood rat; Sylvilagus bach- mani (Waterhouse ) , brush rabbit; S. audubonii (Baird), cottontail rabbit; Canis familiaris L., dog (puppies) ; Anser cincreus, goose; Anas boscas L., duck; Circus rufus (Gm.), hawk; Turdus merida L., blackbird; Ciconia ciconia L., white stork; Vidtur fulvus Briss., tawny vulture; Marmota monax (L.), wood- chuck; Microtus pennstjlvanicus (Ord.), field vole ; Speotyto cunicidaria hypugaea (Bonaparte), western burrowing owl; Molothrus ater (Boddaert), cowbird; common quail or bob-white. Insusceptible species : Frog (cat and opossum reported as "refractory"). Geographical distribution : United States, Canada, Argentina. Induced disease : In horse, initial fever, then signs of fatigue, somnolence; occa- 1254 MANUAL OF DETERMINATIVE BACTERIOLOGY sional excitability followed by incoor- dinatedactionof limbs, disturbed equilib- rium, grinding of teeth, paresis and varied paralyses; frequently inability to swal- low, paralysis of lips and bladder, amauro- sis ; case fatality about 50 per cent ; recovery without sequelae in mild cases ; death within 3 to 8 days in severe cases. In man (childi'en particularly vulnera- ble), a profound, acute, disseminate and focal encephalomj'elitis characterized by intense vascular engorgement, perivascu- lar and parenchymatous cellular infiltra- tion and extreme degenerative changes in the nerve cells. In chick embryo, ex- cessive increase of virus continuing until just before host's death, virus being found eventually throughout the egg but most concentrated in the embryo ; vaccines made from virus grown in chick embryo and then inactivated are especially effec- tive because of the high titer of virus represented in them ; increased resistance with age characteristic of chorioallantoic membrane as well as of hatched chick ; rounded acidophilic masses occur usually near periphery of nucleus in embiyonic nerve cells; no such inclusions are found as a result of infection with Borna disease virus or poliomyelitis virus. Transmission : Experimentally by tick, Dermacenlor andersoni Stiles {IXODl- DAE), passing through eggs to offspring ; this tick is infective to susceptible ani- mals on which it feeds as larva, nymph or adult . Experimentally by Aedes acgypii L. (to guinea pig and horse, preinfective period 4 to 5 days ; insects retain virus for duration of life; not to eggs of infected mosquitoes; not passed from males to females or by males from female to fe- male), A. albopictns, A. atropalpvs, A. cantator, A. dorsalis, A. nigromacrdis, A. sollicilans, A. taeniorhynchus , A. triseria- tus, and A. vexans {CULICIDAE). Triatoma sanguisxiga (Le Conte) {RE- DUVIIDAE) has been found infected in nature and has transmitted virus ex- perimentally to guinea pigs. The Ameri- can dog tick, Dermncentor variabilis Say [IXODIDAE) has been infected by inoculation, not by feeding; it has not been shown to transmit. Serological relationships : Neutralizing antibodies are formed as a result of vac- cination with inactive, formolized virus; antigenicity of formalin-inactivated virus as well as of active virus is blocked in the presence of antiserum. In rabbit, cere- bral resistance is coincident with pres- ence of neutralizing antibody in spinal fluid. In guinea pig, therapy with spe- cific antiserum ineffective if begun after onset of encephalitis ; effective if begun within 24 to 48 hours of peripheral inocu- lation. No cross neutralization reaction with lymphocytic choriomeningitis virus, Japanese B encephalitis virus or St. Louis encephalitis virus. Constituent strains (typical Western and Eastern) do not give cross neutralization reactions, but do show the presence of common antigens by cross reactions in complement fixation not shared with such other viruses as Japanese B encephalitis virus, St. Louis encephalitis virus. West Nile encepha- litis virus, lymphocytic choriomeningitis virus. Sera of human cases may be nega- tive by complement fixation tests a few days after onset, yet strongly strain- specific during second week of illness. Immunological relationships : Young of immunized guinea pigs are immune to homologous strain at least a month after birth. No cross immunity between Western and Eastern strains of equine encephalitis virus. Thermal inactivation : At 60° C, not at 56° C, in 10 minutes. Filterability : Passes collodion mem- branes 66, not 60, millimicrons in average pore diameter. Passes Berkefeld V, N, and W, finest Handler, and Seitz filters. Other properties : Inactivated below pH 5.5. Viable at least a year, dry in vacuum. Particle diameter estimated from filtration experiments to be 20 to 30 millimicrons. Electron micrographs show particles as spherical or disk- shaped, about 39 millimicrons in diameter with round or oval region of high density within each; older preparations show FAMILY ERRONACEAE 1255 comma-shaped particles. Sedimentation constant, mean 265.5 X IQ-i^ ± 5.4 X 10-13 (range 252 to 276 X IQ-") . Specific volume 0.864. Molecular weight of lipo- nucleoprotein complex behaving as the virus calculated as 152 million, approxi- mately 250 particles giving 50 per cent infection; material contains 4 per cent carbohydrate. Absorption of ultraviolet light reaches a peak at about 2600 A., a broad minimum at about 2450 A., and an increase at 2200 A. Strains : The Western strain (so-called Western equine encephalitis virus) may be considered as type of a large group of variants met in nature ; some produce clinically milder disease than others (Birch, Am. Jour. Vet. Res., 2, 1941, 221-226) ; they may change in virulence on passage in experimental hosts. The Eastern strain (so-called Eastern equine encephalitis virus) has been studied ex- tensively also, and has been found to differ from the type strain especially : in more rapid course of induced disease in the horse ; in being experimentally trans- missible to sheep, pig, dog, cat and the European hedgehog; in its localization in eastern coast states and absence from the area between California and Wisconsin, where the type strain is found ; in failure experimentally to infect A'edes aegypti unless inoculated into body cavity by needle punctui'e, whereupon it persists and can be transmitted ; and in failure of cross-neutralization with the western strain. A strain produced by serial passage in pigeons is reported to have caused no obvious reaction in horses but to have induced the formation of neu- tralizing antibodies. A Venezuelan strain differs from the type in comple- ment-fixation reactions; it induces in man a mild disease, characterized by malaise, fever, headache or drowsiness, and uneventful recovery (Casals et al.. Jour. Exp. Med., 77, 1943, 521-.530). Literature: Bang, Jour. Exp. Med., 77, 1943, 337-344; Bauer et al., Proc. Soc. Exp. Biol, and Med., 33, 1935, 378-382; Beard et al.. Science, 87, 1938, 490; Birch, Am. Jour. Vet. Res., 2, 1941, 221-226; Casals and Palacios, Science, 94, 1941, 330; Covell, Proc. Soc. Exp. Biol, and Med., 32, 1934, 51-53; Cox, ibid., 33, 1936, 607-609; Cox and Olitsky, Jour. Exp. Med., 63, 1936, 745-765; ^4, 1936, 217-222, 223-232; Cox et al., U. S. Pub. Health Service, Public Health Rept., 66, 1941, 1905-1906; Davis, Am. Jour. Hyg., 32 (C), 1940, 45-59 ; Eklund and Blumstein, Jour. Am. Med. Assoc, 111, 1938, 1734- 1735; Feemster, Am. Jour. Public Health, 28, 1938, 1403-1410; Finkelstein et al.. Jour. Inf. Dis., 66, 1940, 117-126; Fother- gill and Dingle, Science, 88, 1938, 549- 550; Fothergill et al., New England Jour. Med., 219, 1938, 411 ; Giltner and Shahan, Science, 78, 1933, 63-64; Jour. Am. Vet. Med. Assoc, 88, CN.S. 41), 1936, 363-374; Graham and Levine, Am. Jour. Vet. Res., 2, 1941, 430-435; Grundmann et al.. Jour. Inf. Dis., 72, 1943, 163-171 ; Havens et al., Jour. Exp. Med., 77, 1943, 139-153 ; Higbie and Howitt, Jour. Bact., 29, 1935, 399- 406; Howitt, Jour. Inf. Dis., 55, 1934, 138-149; 61, 1937, 88-95; 67, 1940, 177- 187; Science, 88, 1938, 455-456; Howitt and Van Herick, Jour. Inf. Dis., 71, 1942, 179-191; Kelser, Jour. Am. Vet. Med. Assoc, 82, 1933, 767-771 ; King, Jour. Exp. Med., 71, 1940, 107-112; 76', 1942, 325-334; Kitselman and Grundmann, Kansas Agr. Exp. Sta., Tech. Bull. 50, 1940, 1-15; Merrill and TenBroeck, Jour. Exp. Med., 62, 1935, 687-695; Meyer et al., Science, 74, 1931, 227-228; Mitchell et al., Cana- dian Jour. Ccrmp. Med., 3, 1939, 308-309; Morgan, Jour. Exp. Med., 74, 1941, 115- 132; Morgan et al., ibid., 76, 1942, 357- 369; Olitsky et al., ibid., 77, 1943, 359- 374; Remlinger and Bailly, Compt. rend. Soc. Biol., Paris, 121, 1936, 146-149; 122, 1936, 518-519; 123, 1936, 562-563; Sabin and Olitsky, Proc. Soc. Exp. Biol, and Med., 38, 1938, 597-599; Sellards et al., Am. Jour. Hyg., 33 (B), 1941, 63-68; Shahan and Eichhorn, Am. Jour. Vet. Res., 2, 1941, 218-220; Sharp et al., Proc. Soc. Exp. Biol, and Med., 51, 1942, 206- 207; Arch. Path., 36, 1943, 167-176; Syverton and Berry, ibid., 34, 1936, 822- 1256 MANUAL OF DETERMINATIVE BACTERIOLOGY 824; Jour. Bact., S3, 1937, 60; Am. Jour. Hyg., 82 (B), 1940, 19-23; SS (B), 1941, 37-41; Jour. Exp. Med., 78, 1941, 607- 529; Taylor et al., Jour. Inf. Dis., 67, 1940, 59-66; 69, 1941, 224-231; 73, 1943, 31-41 ; TenBroeck, Arch. Path., 25, 1938, 759 (Abst.); TenBroeck and Merrill, Proc. Soc. Exp. Biol, and Med., 31, 1933, 217-220; TenBroeck et al.. Jour. Exp. Med., 62, 1935, 677-685; Traub and Ten Broeck, Science, 81, 1935, 572; Tyzzer and Sellards, Am. Jour. Hyg., S3 (B), 1941, 69-81; Tyzzer et al.. Science, 88, 1938, 505-506; van Roekel and Clarke, Jour. Am. Vet. Med. Assoc, 94 (N.S. 47), 1939, 466-468; Webster and Wright, Science, 88, 1938, 305-306; Wesselhoeft et al.. Jour. Am. Med. Assoc, 111, 1938, 1735-1740; Wright, Am. Jour. Hyg., 36, 1942, 57-67. 8. Erro bornensis spec. nov. From Borna, name of a town in Saxony where a severe epizootic occurred in 1894 to 1896. Common name : Borna-disease virus. Hosts : Horse, cow, sheep, perhaps deer. Experimentally, also rabbit, guinea pig, rat (more susceptible when old than when younger), mouse; Macaca mvlatta (Zim- mermann), rhesus monkey. Insusceptible species: Ferret, cat, pigeon; probably dog. Geographical distribution : Wurtem- burg, Germany, North and South Amer- ica, Hungary, Russia, Belgium, France, Italjs Roumania. Induced disease: In horse, encephalo- myelitis characterized by lassitude, in- difference to external stimuli; later intermittent excitement, difficulty in mastication and deglutition, spasms in various muscles, champing, excessive salivation ; pupils unequal in size ; paraly- sis of hindquarters, tail, muscles of tongue, or muscles of back; temperature usually normal ; death in 20 to 37 hours or, less often, recovery after about 1 to 3 weeks. Virus may pass placenta and infect fetus in pregnant animals. Transmission : To rabbit, experimen- tally by feeding and by injection in- tracerebrally, intraocularl}'', nasally, intravenously, subcutaneously, or intra- peritoneally ; not by living in same cage. Immunological relationships: No cross immunity conferred by the Western strain of equine encephalomyelitis virus. Isolate of Borna disease virus from the horse immunizes rabbits against isolate from sheep, and vice versa. Herpes and rabies viruses do not immunize rabbits against subsequent infection by Borna disease virus. Thermal inactivation : At 50 to 57° C in 30 minutes; at 70° C in 10 minutes. Filterability : Passes Berkefeld N and Mandler filters, but with difficulty. Passes collodion membranes of average pore diameter 400 millimicrons readily, 200 millimicrons with difficulty, 175 milli- microns not detectibly. May be sep- arated by differential filtration from louping-ill virus, which will pass even a 125-millimicron membrane. Other properties : Particle size esti- mated from filtration data as 85 to 125 millimicrons. Optimum pH for stability in broth at 15 to 20° C is 7.4 to 7.6; very sensitive to greater alkalinity. Viable after 327 days dry at laboratory tempera- tures. Viable at least 6 months in 50 per cent glycerine. Inactivated by putre- faction in 5 days ; by 1 per cent carbolic acid in 4, not in 2, weeks. Literature: Barnard, Brit. Jour. Exp. Path., 14, 1933, 205-206; Covell, Proc. Soc. Exp. Biol, and Med., 32, 1934, 51-53; Elford and Galloway, Brit. Jour. Exp. Path., 14, 1933, 196-205; Howitt and Meyer, Jour. Infect. Dis., 54, 1934, 364- 367; Nicolau and Galloway, Brit. Jour. Exp. Path., 8, 1927, 336-341, and in Medical Research Council, Special Re- port Series No. 121, London, 1928, 90 pp., Ann. Inst. Pasteur, 44, 1930, 673-696; ^5, 1930, 457-523 ;Zwicketal.,Ztschr. Infek- tionskr. parasit. Krankh. u. Hyg. d. Haustierc, 30, 1926, 42-136; 32, 1927, 150-179. FAMILY ERROXACEAE 1257 Genus II. Legio gen. nov. Viruses of the Poliomj-elitis Group, often recoverable from feces of infected hosts probably because of involvement of some part of the alimentary tract ; usually there is also obvious involvement of some part of the nervous system. Generic name from Latin legio, an army or legion. The type species is Legio dehilitans spec. nov. Key to the species of genus Legio. I. Affecting man (see also IV below) . 1. Legio dehilitans. 2. Legio crebea. 3. Legio simvlans. II. Latent in, or affecting, mouse. 4. Legio initris. III. Affecting birds. 5. Legio gallinae. IV. Affecting swine and swineherds. 6. Legio suariorian. 1. Legio dehilitans spec. nov. From Latin debilitare, to weaken or maim. Common names: Poliomyelitis virus, virus of infantile paralysis. Hosts : HOMINIDAE — Homo sapiens L., man. Experimentallj-, Cercopiihecus aethiops sabaeus, green African monkey; Macaca mordax; M. nndatla, the rhesus monkej'; M. irus, the cynomolgus mon- key; mona monkey; for some isolates, Sigmodon hispidus Say and Ord, cotton rat ; mouse ; guinea pig; white rat. Insusceptible species : Sheep ("refrac- tory"biit forms neutralizing antibodies), chicken. Geographical distribution; Almost world-wide. Induced disease : In man, probably sub- clinical in most cases, in view of the presence of specific antibodies in sera from the great majority of adults in all parts of the world; virus probably infects some part of the alimentary tract, being found in stools of most clinical cases, of most apparently healthy contacts, and even of some individuals who have recov- ered from abortive attacks (in one case 123 days after attack); clinical disease, largely in children, is characterized by invasion of central nervous system, with effects ranging from sore throat, fever. vomiting, and headache to sudden and severe paralysis; the muscles most often involved are those of the legs, but there may be paralysis of abdominal or inter- costal muscles. Virus not in urine or saliva, rarely in nasal washings; more of- ten in stools of young than of old patients ; in walls of pharynx, ileum, descending colon. Virus has been recovered from sewage. Incidence and fatality affected by racial characteristics, the first lower and the second higher in negroes than in whites in the L'nited States. In monkey, similar disease, no virus in blood, relapse with reappearance of virus reported; in isolated intestinal loops, infection does not occur through normal mucosa in absence of intestinal contents; disease more severe in summer than in autumn, in autumn than in winter; more severe in older than in younger monkeys; no im- munit}' follows inoculation unless obvious disease occurs. Transmission : Transmission in milk has been suspected and at times con- firmed. Virus has been recovered from mi.xed samples of flies in an epidemic area. No definite arthropod vector has been incriminated. Experimentally, in Cercopithecns aethiops sabaeus, the green 1258 MANUAL OF DETERMINATIVE BACTERIOLOGY African monkey, by intracerebral, intra- nasal, and intraabdominal inoculation. Serological relationships : Specific neu- tralizing antibodies arise after experi- mental infection in monkeys, but rein- fection is not prevented; only a minority of human convalescent sera neutralize virus in vitro, the most potent sera prob- ably being obtained from those with transient or light paralysis. Cross neu- tralization between monkey-passage and murine (cotton-rat and mouse) strains. No cross -neutralization reaction with lymphocytic choriomeningitis virus. Is- olates differ somewhat antigenically, homologous titers being higher than heterologous titers in some neutralization tests. Thermal inactivation : At or below 75° C in 30 minutes. Filterability : Passes membrane about 35, not 30, millimicrons in average pore diameter. Other properties : Infectivity of virus maintained well at —76° C or in glycerine but poorly when dried or just frozen. Inactivated readil.y bj^ hydrogen peroxide. Particle diameter estimated as about 12 millimicrons by filtration studies. Precipitated by half-saturated ammonium sulphate solutions. Electron micrographs show elliptical particles 20 to 30 millimicrons in diameter; impure infectious materials show long threads 20 by 75 to 500 millimicrons in size. Component probably virus has sedimenta- tion constant 820° = 62 X \0~^^ cm per sec. per dyne. Inactivated by potassium hydroxide, copper sulfate and potassium permanganate. Stable from pH 2.2 to 10.4 for 2 hours at 37° C. Literature : Armstrong and Harrison, U. S. Pub. Health Service, Public Health Rept., 50, 1935, 725-730; Aycock, Am. Jour. Hyg., 7, 1927, 791-803; Burnet and Jackson, Austral. Jour. Exp. Biol, and Med. Sci., 17, 1939, 261-270; 18, 1940, 361-366; Burnet et al., ibid., 17, 1939, 253-260, 375-391; Elford et al., Jour. Path, and Bact., I+O, 1935, 135-141 ; Flex- ner, Jour. Exp. Med., 62, 1935, 787-804; 63, 1936, 209-226; 65, 1937, 497-513; Card, ibid., 71, 1940, 779-785; Gordon and Len- nette. Jour. Inf. Dis., 64, 1939, 97-104; Harmon, ibid., 58, 1936, 331-336; Heaslip, Austral. Jour. Exp. Biol, and Med. Sci., 16, 1938, 285-286; Howitt, Jour. Inf. Dis., 51, 1932, 565-573; 53, 1933, 145-156; Hud- son and Lennette, Am. Jour. Hyg., 17, 1933, 581-586; Jungeblut and Bourdillon, Jour. Am. Med. Assoc, 128, 1943, 399- 402; Jungeblut and Sanders, Jour. Exp. Med., 72, 1940, 407-436; 76, 1942, 127-142; Jungeblut et al., ibid., 75, 1942, 611-629; 76, 1942, 31-51; Kessel et al.. Am. Jour. Hyg., 27, 1938, 519-529 ; Jour. Exp. Med., 74, 1941, 601-609 ; Kolmer et al., Jour. Inf. Dis., 61, 1937, 63-68; Kramer et al.. Jour. Exp. Med., 69, 1939, 46-67 ; Lennette and Hudson, Jour. Inf. Dis., 58, 1936, 10-14; Loring and Schwerdt, Jour. Exp. Med., 75, 1942, 395-406 ; McClure and Langmuir, Am. Jour. Hyg., 35, 1942, 285-291 ; Mel- nick, Jour. Exp. Med., 77, 1943, 195-204; Moore and Kessel, Am. Jour. Hyg., 38, 1943, 323-344; Moore et al., ibid., 36, 1942, 247-254; Morales, Jour. Inf. Dis., 46, 1930, 31-35; Olitsky and Cox, Jour, Exp. Med., 63, 1936, 109-125; Paul et al.. Am. Jour. Hyg., 17, 1933, 587-600; 601- 612; Jour. Exp. Med., 71, 1940, 765-777; Sabin, ibid., 69, 1939, 507-516; Sabin and Olitsky, ibid., 68, 1938, 39-61 ; Sabin and Ward, ibid., 73, 1941, 771-793; 74, 1942, 519-529; 75, 1942, 107-117; Sabin et al.. Jour. Bact., 31, 1936, 35-36 (Abst.); Sanders and Jungeblut, Jour. Exp. Med., 75, 1942, 631-649; Schultz and Gebhardt, Jour. Inf. Dis., 70, 1942, 7-50; Schultz and Robinson, ibid., 70, 1942, 193-200; Stimpert and Kessel, Am. Jour. Hyg., 29, (B), 1939, 57-66; Theiler, Medicine, 20, 1941, 443-462; Theiler and Bauer, Jour. Exp. Med., 60, 1934, 767-772 ; Trask and Paul, ibid., 58, 1933, 531-544; 73, 1941 , 453-459 ; Trask et al., ibid., 77, 1943, 531-544 ; Turner and Young, Am. Jour. Hyg., 37, 1943, 67-79; Wolf, Jour. Exp. Med., 76, 1942, 53-72 ; Young and Merrell, Am. Jour. Hyg., 37, 1943, 80-92. FAMILY ERKONACEAE 1259 2. Legio erebea spec. nov. From Latin erebeus, belonging to the Lower World. Common names: Choriomeningitis vi- rus, lymphocytic choriomeningitis virus. Hosts : MURIDAE — Mns muscrdvs L., graj' or white mouse. HOM I N IDAE — Homo sapiens L. , man . CERCOPITHECI- DAE — Macaca mulatla, rhesus monkey. P^xperimentally, also guinea pig; white rat; dog (masked); ferret (masked); Macaca irus, crab-eating macaque; Syri- an hamster; chick- or mouse-embryo serum-Tyrode solution culture; chick embryo. Insusceptible species: Pig, rabbit, field vole, bank vole, canary, hen, para- keet. Geographical distribution: France, England, United States. Induced disease : In white mouse, more virulent in young than in old individuals ; infection may take place in utero or soon after birth; some mice become carriers after recovery, with virus in organs, blood, urine, and nasal secretions; car- riers are immune to large intracerebral inoculations of virus; expei'imentallj^ 5 to 12 days after intracerebral inoculation of susceptible mice, somnolence, photo- phobia, tremors of the legs, tonic spasms of muscles in the hindquarters upon stimulation ; recovery or death. In man, disease may be subclinical at times as shown by the fact that some supposedly normal sera contain specific antibodies; not all clinical cases develop protecting antibodies against testing strains, so that disease may be somewhat commoner than can be ascertained readily ; in all cases benign, but in the more severe of these an acute aseptic meningitis ; after incubation period of !§ to 3 days, spells of fever ex- tending as long as 3 weeks ; late in the disease there may be a meningeal reaction both clinically and cytologically ; lympho- cytes and some large mononuclear cells appear in the meningeal fluids, although symptoms remain benign ; there may be virus in the blood from the beginning of fever to the end of the second week ; the spinal fluid is not infective at first but may become so before there is a change in cell count ; urine and saliva remain unin- fect ious. Transmission : In white mouse, by con- tact with mice infected when young, not with those infected when old; nasal mucosa considered portal of entry. In wild gray mouse of the same species, Mus muscvlus, by contact but less readily than in white mouse. Experimentally, by mosquito, Aedes aegypti L. (C ULICI- DAE), at 26 to 34° C ; by bedbug, Cimex lectularius {CIMIDAE), but defecation on site of bitten area is essential, bite alone being ineffective. Experimentally, to guinea pig, by application of virus to normal and apparently intact skin; not by contamination of food or litter. Serological relationships: Serum of recovered subjects usually neutralizes choriomeningitis virus. Hyperimmune serum is ineffective against pseudo-Iym- phocytic choriomeningitis virus and hy- perimmune serum for that virus is inef- fective in its turn when used with choriomeningitis virus. No cross neu- tralization with St. Louis encephalitis virus. A specific soluble antigen asso- ciated quantitatively with virus in all hosts fixes complement in the presence of immune serum; virus does so poorly if at all; the anti-soluble-substance anti- bodies seem to be independent of virus- neutralizing antibodies. A soluble pro- tein, readily separable from virus, gives a specific precipitin reaction with immune serum ; antibodies concerned are probably not the virus-neutralizing antibodies. Immunological relationships : Intra- peritoneal injection of about 160 intra- cerebral lethal doses has been found to protect the white mouse against infection by subsequent intracerebral injection of 10,000 lethal doses. The immune mouse differs from the immune guinea pig in showing no neutralizing antibodies in its blood; even the guinea pig may develop resistance before antibodies appear in its serum. Formalized vaccines made from 1260 MANUAL OF DETERMINATIVE BACTERIOLOGY guinea pig tissues immunize the guinea pig but vaccines made from mouse tissues do not. Mice immune to this virus are susceptible to infection with pseudo- lymphocytic choriomeningitis virus and vice versa. Thermal inactivation : At 55 to. 56° (J in 20 minutes. Filterability : Passes Berkefeld V, X, and W filters and, with difficulty, a Seitz asbestos pad.- Other properties : Infective at least 206 days in storage at 4 to 10° C in 50 per cent neutral glycerine in 0.85 per cent saline. Infective particle calculated to be 37 to 55 millimicrons in diameter on the basis of centrifugation studies; 40 to 60 milli- microns by ultrafiltration tests. Inac- tivated by soap with loss of mouse-im- munizing capacity. Literature : Armstrong and Dickens, U. S. Pub. Health Service, Public Health Rept., 60, 1935, 831-842; Armstrong and Lillie, ibid., Jfi, 1934, 1019-1027; Arm- strong and Wooley, ihid., 50, 1935, 537- 541; Jour. Am. Med. Assoc, 109, 1937, 410-412; Baird and Rivers, Am. Jour. Pub. Health, 28, 1938, 47-53; Casals- Ariet and Webster, Jour. Exp. Med., 7/, 1940, 147-154; Dalldorf, ibid.. 70, 1939, 19-27; Dalldorf and Douglass, Proc. Soc. Exp. Biol, and Med., 39, 1938, 294-297; Findlay and Stern, Jour. Path, and Bact ., 48, 1936, 327-3.38; Findlay et al.. Lancet, 230, 1936 (/), 650-654; Howard, Jour. Inf. Dis., 64, 1939, 66-77; Laigret and Durand, Compt. rend. Acad. Sci., 203, 1936, 282-284; Lepine and Sautter, Ann. Inst. Pasteur, 61, 1938, 519-526; Lepine et al., ibid., 204, 1937, 1846-1848; Mac- Callum and Findlay, Brit. Jour. Exp. Path., 21, 1940, 110-116; Milzer, Jour. Inf. Dis., 70, 1942, 152-172; Rivers aiid Scott, Jour. Exp. Med., 63, 1936, 415-432 ; Scott and Elford, Brit. Jour. Exp. Path., 20, 1939, 182-188; Scott and Rivers, Jour. Exp. Med., 63, 1936, 397-414; Shaugnessy and Zichis, ibid., 72, 1940, 331-343; Smadel and Wall, ibid., 72, 1940, 389-405 ; 75, 1942, 581-591; Smadel et al., Proc. Soc. Exp. Biol, and Med., 40, 1939, 71-73; Jour. Exp. Med., 70, 1939, 53-66; 7^ 1940, 43-53; Stock and Francis, ibid., 77, 1943, 323-336; Traub, Science, 81, 1935, 298- 299; Jour. Exp. Med., 63, 1936, 533-546, 847-861; 64, 1936, 183-200; 66, 1937, 317- 324; 68, 1938, 9.5-110, 229-250; 69, 1939, 801-817. 3. Legio simulans spec. nov. From Latin simulare, to imitate, in reference to resemblance of this virus to the preceding in many respects, though not in size or antigenic properties. Common name : Pseudo-lymphocytic choriomeningitis virus. Hosts : HOMINIDAE — Ho7no sapiens L., man. Experimentally, also mouse, guinea pig, rhesus monkey; chorioallan- toic membrane of chick embryo. Induced disease : In man, benign asep- tic lymphocj'tic meningitis with virus in cerebro-spinal fluid; severe frontal headache, drowsiness, irritability, vomit- ing, eventual complete recovery. In mouse, experimentally, roughened fur, spontaneous tremor, hunched attitude, irritability, clonic movements ending with tonic convulsions on stimulation, temporary recovery from spasm with survival a few hours or instant death. Serological relationships : Hyperim- mune sera for lymphocytic choriomenin- gitis virus are ineffective for this virus, and vice versa. In man, after recovery, neutralizingantibody is strong at 1 month, fading before 7 months. Immunological relationships: Mice ac- (juire specific resistance to reinfection after experimental disease ; mice immune to lymphocytic choriomeningitis virus are susceptible to pseudo-lymphocytic chori- omeningitis virus and vice versa. Thermal inactivation : At 56° C, not at 45° C, in 30 minutes. Filterability : Passes Berkefeld V, not N, filter candle; Gradacol membrane of 320, not 300, millimicron average pore diameter. Other properties : Particle diameter calculated to be not above 150 to 225 millimicrons, from filtration experiments. FAMILY ERRONACEAE 1261 Viable at least 1 month at 4° C, at least 1 year in 50 per cent glycerine, 40 da3^s in 0.25 per cent phenol, 1 year when dried from frozen material. Inactivated by 0.05 per cent formalin at 4° C in 48 hours ; by boiling in 5 minutes. Literature: MacCallum et al., Brit. Jour. Exp. Path., 20, 1939, 260-269. 4. Legio muris spec. nov. From Latin mus, mouse. Common names : Mouse-poliomyelitis virus, Theiler's-disease virus. Host : MURIDAE — Mus miisculus L., white mouse. Insusceptible species : CERCO- PITHECIDAE—Macaca mulatta (Zim- mermann), rhesus monke3^ Geographical distribution : United States, Japan, Germany, Palestine ; prob- ably widespread wherever white mice are raised. Induced disease : In white mouse, ordinarily no obvious disease, virus oc- curring in feces and not being recoverable from thoracic or abdominal viscera or head (probable source is in abdominal wall ; virus has been recovered most abundantly from intestinal contents, in moderate amounts from walls of intestine and in smaller concentration from mesen- teric lymph glands) ; occasionally, in- dividual mice show flaccid paralysis of hind legs, and brain or spinal-cord sus- pensions from these contain the virus ; mice inoculated intracerebrally show flaccid paralysis in 7 to more than 30 days, first in one limb, later usually in all ; the tail does not become paralyzed; very ■young inoculated mice maj' die without first showing paralysis ; very old inocu- lated mice may become infected without showing obvious disease ; some affected mice recover and those showing residual paralysis may become carriers of virus. In affected, experimentally inoculated mice, acute necrosis of ganglion cells of anterior horn of spinal cord ; necrosis also of isolated ganglion cells of cerebrum. Later, marked neuronophagia. Perivas- cular infiltration in brain and spinal cord. The reciprocal of the incubation period has been found approximatelj' propor- tional to the logarithm of the amount of virus inoculated, thus serving to measure the concentration of samples of virus. Old mice less susceptible than young. Transmission : Experimentally, by in- tracerebral, intranasal and intraperi- toneal inoculation. Has been found to persist in adult flies, Musca domestica L. (M use I DAE) and other species, as long as 12 days after experimental feed- ing whereas mouse-adapted human polio- myelitis virus persists only 2 days in Musca domestica and not at all in some other species. Serological relationships : Sera con- taining antibodies to the Lansing strain of human poliomj'elitis virus fail to pro- tect against mouse poliomyelitis virus. Immunological relationships : Recov- ered mice are immune to various hetero- logous isolates or strains. No evidence of immunological relationship with virus of human poliomyelitis has been obtained, save that mice paralyzed with mouse poliomyelitis virus show some resistance to infection with the Lansing strain of hu- man poliomyelitis virus ; this has been interpreted as possibly no more than an interference phenomenon, since it seems to depend on actual paralysis. Filterability : Passes Berkefeld X and other Berkefeld filters and Chamberland L3 filter. Other properties : Viable at least 14 months at —78° C ; at least 150 days in 50 per cent glycerine at 2 to 4° C. Most stable near pH 8.0 and pH 3.3. Inac- tivated readily at 37° C by 1 per cent hydrogen peroxide. Particle diameter estimated as 9 to 13 millimicrons from filtration studies. Sedimentation con- stant, Sjo" = 160 to 170 X 10-1' cm per sec. per dyne. Literature : Bang and Glaser, Am. Jour. Hj'g., 37, 1943, 320-324; Gahagan and Stevenson, Jour. Inf. Dis., 69, 1941, 232- 237; Gard, Jour. Exp. Med., 72, 1940, 69-77; Gard and Pedersen, Science, 94, 1941, 493-494; Gildemeister and Ahlfeld, 1262 MANUAL OF DETERMINATIVE BACTERIOLOGY Cent. f. Bakt., I Abt., Orig., U2, 1938 , 144-148; Iguchi, Kitasato Arch. Exp. Med., 16, 1939, 5&-78; Olitsky, Jour. Exp. Med., 72, 1940, 113-127; Theiler, Science, 80, 1934, 122; Jour. Exp. Med., 65, 1937, 705-719 ; Theiler and Gard, ihid., 72, 1940, 49-67, 79-90; Young and Cum- berland, Am. Jour. Hyg., 37, 1943, 216- 224. 5. Legio gq.llinae spec. nov. From Latin gallina, hen. Common names : Avian encephalomye- litis virus, infectious avian encephalo- myelitis virus. Host: PHASIANIDAE—Gallus gal- lus (L.), chicken (embryo not suscepti- ble ; in culture media, minced whole embryo in serum-Tyrode solution suffices to maintain virus, but embryo brain alone does not ) . Insusceptible species : All tested spe- cies other than birds. Geographical distribution : United States. Induced disease : In chicken, fine or coarse tremors of whole body or only of head and neck or of legs; progressive ataxia; eyes dull, some loss of weight, weakness of legs, and progressive inco- ordination of leg muscles ; somnolence precedes death; about 75 per cent die within 5 days of onset, 90 per cent within a week, the remainder showing a stag- gering, ataxic gait for weeks, some con- tinuously tremulous; recovered birds, however, may produce eggs well ; micro- scopic focal collections of glia cells, perivascular infiltration, degeneration of Purkinje 's cells and degeneration of nerve cells ; foci of infiltration throughout brain and spinal cord ; virus not detected in the blood of affected chickens. Transmission : Not through egg. Ex- perimentally, by intracerebral injection. Serological relationships : Specific anti- serum neutralizes homologous virus but not the Eastern strain of equine ence- phalitis virus ; antiserum specific for the latter does not neutralize avian ence- phalomyelitis virus. Filterability : Passes Berk (fdd V and N as well as Seitz 1 and 2 filters; also membranes 73 millimicrons in average pore diameter. Other properties : Survives in 50 per cent glycerine for at least 88 days and frozen for at least 68 days. Infective particle estimated to be 20 to 30 milli- microns in diameter, by filtration studies. Literature : Jones, Science, 76, 1932, 331-332; Jour. Exp. Med., 59, 1934, 781- 798; Kligler and Olitsky, Proc. Soc. Exp. Biol, and Med., 4^, 1940, 680-683 ; Olitsky, Jour. Exp. Med., 70, 1939, 565-582; Olit- sky and Bauer, Proc. Soc. Exp. Biol, and Med., 4^, 1939, 634-636 ; Van Roekel et al., Jour. Am. Vet. Med. Assoc, 93 (N.S. 46), 1938, 372-375. 6. Legio suariorum spec. nov. From Latin suarius, swineherd. Common name : Swineherds'-disease virus. Hosts: SUIDAE — Sus scrofa L., swine. HOMIN IDAE — Homo sapiens L., man. Experimentally, with fever as only symptom, white rat, cat, ferret, mouse ; perhaps Macaca mulatta (Zim- mermann), rhesus monkey. Geographical distribution : Europe. Induced disease : In man, a benign meningitis without sequelae, somewhat similar to lymphocytic choriomeningitis in man; cell counts in spinal fluids may be as high as 1200 to 1400; 4 to 7 (average 8) days after infection, fever lasting 3 to 21 days (average 9); sometimes conjunc- tivitis, more often a reddish maculo- papillose eruption; severe sweating frequent; hemorrhagic tendency; blood in feces; recovery. Blood, urine, feces infectious, not spinal fluid or mucous excretions. Especially affecting young men, not often old men or women, among those having contact with swine or swine- producing quarters. Transmission : Excreta of pigs, even as used for manure, are infective. Experi- mentally, to man, by subdermal or intra- muscular injection. FAMILY ERRONACEAE 1263 * Serological relationships : Serum from recovered cases neutralizes the virus. Immunological relationships : Specific immunity follows attack of the disease. Filterability : Passes Chamberland L2 filter. Literature :Durandetal., Compt. rend. Acad. Sci., Paris, 203, 1936, 830-832, 957- 959, 1032-1034; Arch. Inst. Pasteur de Tunis, 26, 1937, 213-227; 228-249; 27, 1938, 7-17. Genus III. Formido gen. nov. Viruses of the Rabies Group, inducing diseases characterized by involvement of the nervous system only. Generic name from Latin /ornnV/o, a frightful thing. The type and only recognized species is Formido inexorabilis spec. nov. 1. Formido inexorabilis spec. nor. From Latin inexorabilis, implacable. Common name : Rabies virus. Hosts: CAN I DAE — Canis familiaris L., dog. FELIDAE—Felis catus L., do- mestic cat ; F. negripes, black-footed cat ; F. ocreata, wild cat. HOMINIDAE— Homo sapiens L., man. MUSTELI- DAE — Icionyx orangiae, polecat. SCI- URIDAE — Geosciurus capensis, ground squirrel. V IVERRIDAE—Cynictis penicillata, yellow mongoose (yellow meercat) ; Genetta felina (Thunb.), genet cat; Myonax pulverulentus , small, grey mongoose; Suricata suricatta, Cape suri- cate or common meercat. Cattle, sheep, pig, horse , wolf. Cynalopex chama, silver jackal. Phyllostoma superciliatum, vam- pire bat; Desmodus rufus, vampire bat; Artibens planirostris trinitatis, fruit-eat- ing bat. Experimentally, also Mus mus- culus L., white mouse ; Peromyscus polio- notus polionotus (Wagner), white-footed mouse; tissue cultures of 5 or 6-day -old rat- or mouse-embryo brain; chick em- bryo (allantois not regularlj^ infected, but virus regularly reaches brain of em- bryo without injuring it ; chick may hatch with titer of 1:100 or 1:1000 in brain). Chicken; mouse hawk (iJuieo vulgaris); pi- geon, owl, goose; stork (Ciconia ciconia); pheasant {Diardigallus diardi B.P.). Insusceptible species : Reptiles, fish. No mammal is known to be insusceptible. Geographical distribution : Almost world-wide ; absent only from relatively isolated countries or communities. Induced disease : In dog, after a short incubation period (generally less than 10 daj's) altered behavior, hiding, lack of obedience, perverted appetite leading to ingestion of straw, paper, earth, and other unaccustomed materials ; excitement, un- provoked biting (which may transmit the virus to new hosts), aimless wandering, excess salivation, progressive inability to swallow, alteration of bark to characteris- tic high pitched tone; staggering, paresis of hindquarters tending toward paralysis and involvement of anterior parts of the body; paralysis of lower jaw, muscular spasms, marked emaciation, death except perhaps in rare instances. In man, after a relatively long incubation period de- pending on site of implantation (perhaps 27 to 64 days), a uniformly fatal disease, characterized by altered behavior, in- creased excitability, thirst, pharyngeal spasm with progressive inability to swal- low, labored and noisy respiration, death in 3 or 4 days after onset, with or without paroxysm. In sheep, increased sexual desire ; tendency to pull wool from other sheep or themselves; light butting, in- creasing until some ewes, after violent exercise, appear to faint; prostration within 1 to 4 days ; death within 2 days from onset of locomotory paralysis. In mouse, experimentally, by intracerebral inoculation, apathy, sluggishness, rough- ening of hair, tremor, convulsions, pros- tration, death; sometimes flaccid paralysis of hind legs before death. Transmission: Usually by bite of dog or some closely related animal ; occa- sionally by bites of cats ; rarely by bites 1264 MANUAL OF DETERMINATIVE BACTERIOLOGY of rabid horses or cattle. Not by con- tamination of food. In Brazil and Trini- dad, probably by the vampire bat, which has been found infected in nature. Serological relationships: Specific floc- culation of rabies virus occurs in the presence of immune serum from rabbit or guinea pig; strains differ in relative amounts of antigenic constituents, as shown by absorption tests. Comple- ment fixation occurs in the presence of virus and guinea-pig antiserum. Neu- tralizing antibodies are specific. Immunological relationships : Virus ex- posed to ultraviolet light tends to lose its virulence before its immunizing potenc}^ Passive immunization succeeds in white mice if antiserum is injected intracere- brally I hour before, but not 24 hours be- fore or 2 hours after, virus. Chloroform- treated vaccines more effective than phenolized vaccines, but irritative. Thermal inactivation : At 60 to 70° C in 15 minutes; in brain tissues, at 45" C in 24 hours. Filterability : Passes Berkefeld V filter. Other properties : Viable at least 2 months at 5° C in liquid or dry state. In- fective particle between 100 and 240 milli- microns in diameter, by filtration studies. Literature : Bernkopf and Kligler, Brit. Jour. Exp. Path., 18, 1937,481-485 ; Casals, Jour. Exp. Med., 72, 1940, 445-451, 453- 461 ; Covell and Danks, Am. Jour. Path., 8, 1932, 557-572; Dawson, Science, 89, 1939, 300-301 ; Am. Jour. Path., 17, 1941, 177-188; Galloway, Brit. Jour. Exp. Path., 15, 1934, 97-105 ; Goodpasture, Am. Jour. Hyg., 1, 1925, 547-582; Haupt and Rehaag, Ztschr. f. Infektionskrankh., 22, 1921, 76-88, 104-127; Havens and May- field, Jour. Inf. Dis., 50, 1932, 367-376 ; 51 , 1932, 511-518; 52, 1933, 364-373; Hender- son, Vet. Med., 37, 1942, 88-89; Hodes ct al., Jour. Exp. Med., 72, 1940, 437-444; Hoyt et al., Jour. Inf. Dis., 59, 1936, 152- 158; Hurst and Pawan, Lancet, 221, 1931 (2), 622-628; Jour. Path, and Bact., 35, 1932, 301-321 ; Johnson and Leach, Am. Jour. Hyg., 82 (B), 1940, 38-45; Kligler and Bernkopf, Proc. Soc. Exp. Biol, and Med., 39, 1938, 212-214; Am. Jour. Hyg., SS (B), 1941, 1-8; Leach and Johnson, ibid., 32 (B), 1940, 74-79 ; Metivier, Jour. Comp. Path, and Therap., 48, 1935, 245- 260; Peragallo, Giorn. di batteriol. e im- munoL, 18, 1937, 289-290; Snyman, On- derstepoort Jour. Vet. Sci. and Anim. Indust., 15, 1940, 9-140; Webster, Am. Jour. Pub. Health, 26, 1936, 1207-1210; Jour. Exp. Med., 70, 1939, 87-106; Am. Jour. Hyg., 30 (B), 1939, 113-134; Web- ster and Casals, Jour. Exp. Med., 71, 1940, 719-730; 73, 1941, 601-615; 76, 1942, 185-194; Webster and Clow, ibid., 66, 1937, 125-131; Wyckoff, Am. Jour. Vet. Res., 2, 1941, 84-90. Note: The Negri body, a character- istic cell-inclusion in rabies, has been given the following names under the sup- position that it represents stages in the life cycle of a protozoan parasite responsi- ble for the disease: Neuronjctes hydropho- biae by Calkins, Jour. Cutaneous Diseases including Syphilis, 25, 1907, 510; Encepha- litozoon rahiei by Manouelian and Viala, Ann. Inst. Pasteur, 38, 1924, 258; and Glugea lyssae by Levaditi,- Nicolau and Schoen, Ann. Inst. Pasteur, 40, 1926, 1048. FAMILY CHARONACEAE 1265 FAMILY IV. CHARONACEAE FAM. NOV. Viruses of the Yellow-Fever Group, inducing diseases mainlj' characterized b}' fever and necrosis of tissues in the absence of obvious macule, papule, or vesicle formation or of conspicuous involvement of nerve cells. Key to the genera of family Charonaceae. I. Viruses of the Typical Yellow-Fever Group. Genus I. Charon, p. 1265. II. Viruses of the Influenza Group. Genus II. Tarpeia, p. 1268. III. Viruses of the Hog-Cholera Group. Genus III. Tortor, p. 1275. Genus I. Charon gen. nov. Viruses of the Typical Yellow-Fever Group, inducing diseases mainly characterized by acute non-contagious fever. Vectors dipterous insects, so far as known. CJeneric name from Latin Charon, ferryman of the Lower World. The type species is Charon evagatus spec. nov. Key to the species of germs Charon. I. Vectors mosquitoes. 1. Charon evagatus. II. Vectors unknown, perhaps mosquitoes. 2. Charon vallis. 1. Charon evagatus spec. nov. From Latin evagor, to spread abroad. Common name : Yellow-fever virus. Hosts : HOMINIDAE—Homo sapiens L., man. Experimentally, also Cerco- pithecus tantalus Ogilby; C. aethiops, African guenon (symptomless) ; Cerco- cebus torquatus (Kerr), collared manga- bey; Mus musculus L., mouse; Microtus agrestis, field vole; Sciurus vulgaris L., red squirrel; Macaca mulatta (Zimmer- mann), rhesus monkey; Macacus sinicus Indian crown monkey ; M. cynonwlgus ; M. speciosus ; Erinaceus europaeus , hedge hog; Gallus gallus (L.), chicken (tol- erant); Dasyprocta aguti, agouti (serial passage fails). Insusceptible species : Cat, ferret, rab- bit, rat; Cricetus auratus, golden ham- ster; Apodemus sylvaticus, wood vole; Evotomys glareolus, bank vole ; pigeon, canary, pipistrelle bat ; Cricetomys gam- bianus, pouched rat; dog, goat. Geographical distribution : Tropical re- gions in general, especially Central and South America, West Indies, West Africa; anti-mosquito campaigns have tended to eradicate yellow-fever virus from parts of its former range. Induced disease : In man, mild cases may occur, especially in natives where the disease is endemic, but in Europeans generally sudden fever without marked change in pulse rate after a 3 to 6-day incubation period; severe frontal head- ache, pains in the loin and legs and epi- gastric pain; gradual decrease in tem- perature to 98 or 99° F, weakening of pulse and slowing of heart beat in the absence of further temperature changes; jaundice, especially in sclerae, often in skin; albumen in urine, later bile -pig- ments also present ; hemorrhages frequent especially in alimentary canal ; fatty and necrotic changes in the liver; acute degeneration of renal parenchyma, splenic congestion ; death may occur in the early acute state, but is more likelj' about the 1266 MANUAL OF DETERMINATIVE BACTERIOLOGY fifth or sixth day; relapses may occur until 2 or 3 weeks after onset ; case mor- tality varies from 10 to 90 per cent in different epidemics. A transitory im- munity due to transfer of serum anti- bodies through the placenta protects offspring of immune mothers for a short time. Transmission: By mosquitoes, Aedes aegypti L., Aedes leucocelaemxs (D. and S.), Haemogogvs capricorni Lutz (CULICIDAE). The mosquito Aedes aegypti becomes infective, after feeding on a suitable virus source, in 4 days at 37° C, 5 days at 36° C, 6 days at 31° C, 8 days at 25.1° C, 9 to 11 days at 23.4° C, 18 days at 21° C, and 36, not 30, days at 18° C; virus in head, thorax, and ab- domen before bites are infective ; no evidence of transmission of virus through eggs to offspring or to larvae eating in- fected adults. Experimentally, also by Aedes scapidaris (Rondani), A.fluviatilis (Lutz), A. Ivteocephalus , A. apico-an- nulatus {CULICIDAE). Experimen- tally, by feeding, to Macacn mulatta and Cercopithecus aethiops; by rubbing in- fected blood into intact and unshaved skin of monkeys. Serological relationships : Complement - fixation and precipitating antibodies arc specific. Immunological relationships : A specific immunity develops after an attack of the disease or after vaccination with virus grown in media containing tissues of chick embryo minus head and spinal cord. Thermal inactivation : At 55 to 60° C, not at 50° C, in 10 minutes. Filterability : Passes membranes of 55, and to some extent membranes of 50. millimicron average j)ore diameter. Passes Berkefeld V and N, as well as Chamberland F, filters. Other properties : Particle estimated from filtration data to have a diameter of 17 to 28 millimicrons ; by ultracentrifuga- tion data, 19 millimicrons. Inactivated or inhibited by 30-minute exposure to 1:15 formalin, 1:6 ethyl alcohol; 1 :3(X) yellowish eosin, 1 : 50 sodium oleate, 1 :200 liquor cresolis compositus; viable after 30-minute exposure at 30° C to 1:7500 mercuric chloride, 1:150 phenol, 1:1500 hexylresorcinol, 1:150 sodium oleate. Sedimentation constant between 18 and 30 X 10~" cm per sec. per dyne. Viable in 50 per cent glycerine at 2 to 4° C for 58, not for 100, days ; in mouse brain at — 8° C for 160 days. Viability may be lost on simple drying but retained if drying is carried on in vacuo over a desiccating agent. Strains : Distinctive strains have been isolated. One, to which much study has been given, differs from the typical vis- cerotropic strain by possessing marked neurotropic or pantropic characteristics. Literature : Bauer and Hughes, Am. Jour. Hyg., 21, 1935, 101-110; Bauer and Mahaffy, ibid., 12, 1930, 155-174; 175- 195; Bugher and Gast-Galvis, ihid., 39, 1944, 58-66; Bugher et al., ibid., 39, 1944, 16-51; Davis, ibid., 16, 1932, 163-176; Davis and Shannon, ibid., 11, 1930, 335- 344; Davis et al., Jour. Exp. Med., 58, 1933, 211-226; Findlay, Jour. Path, and Bact., 38, 1934, 1-6; Lancet, 227, 1934 (2), 983-985; Findlay and Clarke, Jour. Path, and Bact., 40, 1935, 55-64 ; Findlay and MacCallum, Brit. Jour. Exp. Path., 19, 1938, 384-388; Jour. Path, and Bact., 49, 1939, 53-61; Findlay and Mackenzie, ibid., 43, 1936, 205-208; Findlay and Stern, ibid., 40, 1935, 311-318; Fox and Cabral, Am. Jour. Hyg., 37, 1943, 93-120; Frobisher, Am. Jour. Hyg., 11, 1930, 300- 320; 13, 1931, 585-613; 14, 1931, 147-148; 18, 1933, 354-374; Goodpasture, Am. Jour. Path., 8, 1932, 137-150; Haagen, Deutsch. med. Wochnschr., 60, 1934, 983-988; Hudson, Am. Jour. Path., 4, 1928, 395-430; Klotz and Simpson, ibid., 3, 1927, 483-488; Laemmert and Mous- satche. Jour. Inf. Dis., 72, 1943, 228-231 ; Lloyd et al., Am. Jour. Hyg., 18, 1933, 323-344 ; Trans. Roy. Soc. Trop. Med. and Hyg., 29, 1936, 481-529; Mahaffy et al., Am. Jour. Hyg., 18, 1933, 618-628; Pickels and Bauer, Jour. Exp. Med., 71, 1940, FAMILY CHAROXACEAE 1267 703-717; Ramsey, Am. Jour. Hyg., 13, 1931, 129-163; Sawyer, ibid., 25, 1937, 221-231; Shannon et al.. Science, 88 1938, 110-111; Smith and Theiler, Jour. Exp. Med., 65, 1937, 801-808 ; Smith et al.. Am. Jour. Trop. Med., 18, 1938, 437-468; Soper and De Andrade, Am. Jour. Hyg., 18, 1933, 588-617; Soper et al., ibid., 18, 1933, 555-587; 19, 1934, 549-566; 27, 1938, 351-363; Stefanopoulo and Wassermann, Bull. Soc. Path. Exot., 26, 1933, 557-559; Stokes et al.. Am. Jour. Trop. Med., 8 1928, 103-164; Theiler, Ann. Trop. Med. and Parasit., 24, 1930, 249-272; Theiler and Smith, Jour. Exp. Med., 65, 1937, 767-786, 787-800; Whitman, ibid., 66, 1937, 133-143. 2. Charon vallis spec. nov. From La- tin vallis, valley. Common name : Rift Vallej' fever virus. Hosts : HOMINIDAE — Homo sapiens L., man. BOVIDAE — Bos taurus L., cow; Ovis aries L., sheep; Capra hircus L., goat. Experimentally, also Sciiirus carolinensis, grey squirrel; ferret; Crice- tus auratus, golden hamster; Apodemus sylvaticus, wood mouse; Microtus agrestis field vole ; Muscardinus avellanarius , dor- mo "^se ; rat ; mouse ; Macaca midatta ; M. irus; Cebus fatuellus; C. chrysopus; Hapale jacchus; H. penicillata ; Cerco- pithecus callitrichus (symptomless) ; Ery- throcebus patas (symptomless); Cerco- cebus fuliginosus (symptomless); chick embryo in Tyrode's solution ; chorioallan- toic membrane of chick embrj'O. Insusceptible species : Horse, pig. Geographical distribution : British East Africa. Induced disease : In man, benign dis- ease; after 5§ to 6 days, rigors, pains in back, fever for 12 to 36 hours, followed by recovery, with persistence of acquired immune bodies as long as 4 to 5 years after infection. In sheep (lambs), dull- ness, rapid respiration, collapse and death in a few hours or a chronic course ; focal necrosis in liver. In chorioallantoic membrane of chick embryo, experimen- tally, areas of hyperplasia and of necrosis ; connective tissue inflamed nearby; liver of embryo mottled with necrotic areas. Transmission: Not by contacts. Mos- quito, Taeniorhynchusbrevipalpis {CUL- ICIDAE), suspected as possible vector. Serological relationships: Antisera for psittacosis, dengue fever, and sandfly fever viruses fail to protect against infec- tion with Rift Valley fever virus. Spe- cific neutralizing antibody in intraperi- toneally neutral mixture with Rift Valley fever virus may be dissociated so as to free virus by direct dilution in saline solutions, by intranasal inoculation, or by employment of a small dose, all methods probably imph'ing a dilution effect. Immunological relationships : No cross immunity with yellow-fever or dengue- fever viruses. If Rift Valley fever virus is inoculated into rhesus monkey simul- taneously with yellow-fever virus, the animal tends to be protected against death from yellow fever (interference effect), but one-day earlier inoculation of Rift Valley fever virus does not protect. Thermal inactivation : At 56° C in 40, not 20, minutes. Filterability : Passes Berkefeld V, N, and W filters ; passes Chamberland L2, L3, Lo, L7, Lu and occasionally L13 filters ; passes membranes 150 millimicrons in average pore diameter freely, 90 milli- microns with difficulty, 70 millimicrons not at all. Other properties : Mable at least 8 months at 4° C, more than 4 weeks dry in liver tissues, 6 months in 5 per cent carbolic acid at 4° C. Diameter of in- fective particle estimated from filtration studies to be between 23 and 35 milli- microns. Strains: A neurotropic strain im- munizes lambs without producing obvious illness, if given subcutaneoush'. Literature: Broom and Findlay, Brit. Jour. Exp. Path., U, 1933, 179-181; Daubney et al.. Jour. Path, and Bact., 34, 1931, 545-579; Findlay, Trans. Roy. 1268 MANUAL OF DETERMINATIVE BACTERIOLOGY Soc. Trop. Med. and Hyg., 25, 1932, 229- 266; 26, 1932, 157-160; 161-168; Brit. Jour. Exp. Path., 17, 1936, 89-104; Find- lay and MacCallum, Jour. Path, and Bact., U, 1937, 405-424; Findlay and Mackenzie, Brit. Jour. Exp. Path., 17, 1936, 441-447; Findlay et al., ibid., 17, 1936, 431-441; Francis and Magill, Jour. Exp. Med., 62, 1935, 433-448; Horning and Findlay, Jour. Roy. Micr. Soc, 5J^, 1934, 9-17; Mackenzie, Jour. Path, and Bact., 37, 1933, 75-79; 40, 1935, 65-73; Mackenzie et nl., Brit. Jour. Exp. Path., 17, 1936, 352-361; Saddington, Proc. Soc. Exp. Biol, and Med., 31, 1934, 693-694; Schwentker and Rivers, Jour. Exp. Med., 59, 1934, 305-313. Genus II. Tarpeia gen. nov. Viruses of the Influenza Group, inducing diseases characterized principally by involvement of the respiratory tract. Generic name from Latin Tarpeia, name of a Roman maiden who treacherously opened a citadel to an enemy. The type species is Tarpeia alpha spec. nov. Key to the species of genus Tarpeia. I. Infecting man principally. 1. Tarpeia alpha. 2. Tarpeia beta. 3. Tarpeia premens. II. Affecting feline species. 4. Tarpeia felis. III. Affecting domestic cattle (calves). 5. Tarpeia vitulae. IV. Affecting canine species. 6. Tarpeia cants. 7. Tarpeia vulpis. V. Affecting ferrets. 8. Tarpeia viverrae. VI. Affecting domestic fowl. 9. Tarpeia avium. 1 . Tarpeia alpha spec. nov. From first letter of Greek alphabet. Common name : Influenza A virus ; swine filtrate -disease virus. Hosts : HOMINIDAE—Homo sapiens L., man. SUIDAE—Sus .scrofa L., do- mestic swine. Experimentally, also fer- ret, mouse, Macacus irus, hedgehog, rabbit (inapparent infection), guinea pig (inapparent infection), rat (inapparent infection); Mustela sibirica Milne-Ed- wards, Chinese mink; Sciurotarnius da- vidianus Milne-Edwards, David's squir- rel ; chick embryo (some strains produce visible lesions at 36.5° C on chorioallan- loic membrane); minced chick embrj^o in Tyrode's solution. Insusceptible species : Callosciurus caniceps canigenus Howell, Chekiang squirrel ; Eutamias asiaticus senescens Miller, chipmunk. Geographical distribution : World-wide. Induced disease : In man, headache, dizziness, with shivering and muscular pains; rise of temperature on the second day, sometimes with fall on the third and elevation again later; often complicated by bronchitis and bronchopneumonia; I'.emorrhagic and edematous lobular con- sclidation in lungs; virus most easily recoverable from nasopharyngeal wash- ings, but also from nasal secretions and lungs. In swine, virus alone produces only a mild malady (filtrate disease); FAMILY CHARONACEAE 1269 in the presence of Haemophilus influenzae suis a severe maladj' occurs under both natural and experimental conditions; it involves fever, cough, and prostration; many infected animals die. Lungworms, Metasirongylus elongatus and Choero- strongylus pudendotectus {META- STRONGYLIDAE), from infected swine harbor virus at least 2 years, living nieantime in earthworms, such as Allolo- bophora caliginosa (LUM B RIGID AE), which are eaten eventually by swine. The swine are refractory to viral infection during May, June, July, and August, but the disease may be invoked later by successive intramuscular injections of Haemophilus influenzae suis or other stimuli, such as feeding embryonated Ascaris ova. In infected swine, virus oc- curs in turbinates, tracheal exudates, and lungs; not in spleen, liver, kidnej', mesenteric lymph nodes, brain, blood, or mucosa of colon. Xeutralizingantibodies appear later (7th to 10th day) in the mild filtrate disease than in typical swine in- fluenza, in which they appear about the 6th to the 7th day; maximum titer on 14th to 27th daj^ Experimentally in mouse, not contagious as in swine and not dependent on the coexistence of a bacterial component ; death of epithelium of respiratory and terminal bronchioles, complete epithelial desquamation, dilata- tion of bronchioles, collapse of alveoli ; in healing, widespread epithelial prolifera- tion. Experimentally in ferret, mod- erate apathy, lack of appetite, pallor of nose, variable catarrhal symptoms; at acute stage of disease, necrosis of respira- torj^ epithelium of nasal mucous mem- brane, with desquamation of superficial cells, exudation into air passages and inflammatory reaction in the submucosa ; repair follows, beginning on the 6th daj* after infection and becoming essentially complete at the end of 1 month; after recovery, the ferret is immune for 3 months or more, with subsequent waning of resistance ; subsequent subcutaneous inoculations of virus restore immunity. Transmission : Presumably by drop- lets ; for example between cages of ferrets as close as 5 feet apart, even to levels 3 feet higher than cage of diseased individ- uals. Experimentally, from washings of human throats to ferret, mouse, chick embryo (by amniotic route and to allan- toic membrane) ; in mice, bj^ contact and by inhalation of fine droplets. Serological relationships : Neutralizing antibodies common in human sera from individuals above 10 j'ears of age ; rarer in sera from j'oung children ; strongly ef- fective for homologous, weak for hetero- logous, virus in convalescent sera. Solu- ble complement-fixing antigen of swine strain has components in common with antigens of human strains (PR8 and WS). Complement fixation best 10 to 14 days after onset in man. Inactivating capa- city of nasal secretions proportional to level of neutralizing antibodies in blood. Agglutination of red cells by influenza virus is inhibited quantitatively bj' specific antiserum. Immunological relationships: Specific immunization of ferrets, without obvious disease, occurs as a result of intranasal inoculation of egg-passage influenza virus that is not transmissible from ferret to ferret. In mice, immunizing dose is directly proportional to degree of induced immunity; immunity to the strain used in immunization is more effective in gen- eral than that to heterologous isolates of the virus. Filterability : Passes Berkefeld V filter. Other properties: Particle size esti- mated as 80 to 120 millimicrons by filtra- tion studies; 80 to 99 millimicrons by ultracentrifugation (820° = 724 X IQ-i" cm per sec. per dyne); electron micro- graphs show bean or kidney-shaped particles, or round particles with central dense spot , averaging 77.6 millimicrons in diameter. Inactivated by oleic, linolic and linolenic acids without loss of im- munizing ability. Inactivated by ultra- violet radiation. Literature : Andrewes and Glover, 1270 MANUAL OF DETERMINATIVE BACTERIOLOGY Brit. Jour. Exp. Path., 22, 1941, 91-97; Andrewes et al., ibid., 16, 1935, 566-582; Burnet, ibid., 17, 1936, 282-293; 18, 1937, 37-43; 21, 1940, 147-153; Austral. Jour. Exp. Biol, and Med. Sci., 14, 1936, 241-246; 19, 1941, 39-44, 281-290; Eaton, Jour. Bact., 39, 1940, 229-241 ; Eaton and Pearson, Jour. Exp. Med., 72, 1940, 635-643; P^aton and Rickard, Am. Jour. Hyg., 33, (B), 1941, 23-35; Elford et al., Brit. Jour. Exp. Path., 17, 1936, 51-53; Francis, Science, 80, 1934, 457-459 ; Jour. Exp. Med., 69, 1939, 283-300; Francis and Magill, Science, 82, 1935, 353-354; Brit. Jour. Exp. Path., 19, 1938, 284-293; Francis and Shope, Jour. Exp. Med., 63, 1936, 645-653; Francis and Stuart -Harris, ibid., 68, 1938, 789-802; Francis et al., Am. Jour. Hyg., 37, 1943, 294-300; Hirst, Jour. Exp. Med., 75, 1942, 49-64; Hirst et al., ibid., 75, 1942, 495-511 ; Proc. Soc. Exp. Biol, and Med., 50, 1942, 129-133; Horsfall and Lennette, Jour. Exp. Med., 73, 1941, 327-333 ; Hudson et al., ibid., 77, 1943, 467-471 ; Hyde, Am. Jour. Hyg., 36 1942, 338-353; Lennette and Horsfall, Jour. Exp. Med., 73, 1941, 581-599; Loosli et al.. Jour. Inf. Dis., 72, 1943, 142-153; Lush and Burnet, Austral. Jour. Exp. Biol, and Med. Sci., 15, 1937, 375-383; Magill and Francis, Brit. Jour. E.xp. Path., 19, 1938, 273-284; Nigg et al.. Am. Jour. Hyg., 34 (B), 1941, 138-147; Orcutt and Shope, Jour. Exp. Med., 62, 1935, 823-826; Rosenbusch and Shope, ibid., 69, 1939, 499-505; Shope, ibid., 59, 19'34, 201-211 ; 62, 1935, 561-572; 64, 1936, 47-61; 67, 1938, 739-748; 74, 1941, 41-47, 49-68; 77, 1943, 111-126, 127-138; Shope and Francis, ibid., 64, 1936, 791-801; Smillie, Am. Jour. Hyg., 11, 1930, 392- 398; Smith et al.. Lancet, 225, 1933 (2), 66-68; Brit. Jour. Exp. Path., 16, 1935, 291-302 ; Smorodintseff and Ostrovskaya, Jour. Path, and Bact., 44, 1937, 559-566; Stock and Francis, Jour. Exp. Med., 71, 1940, 661-681; Straub, Jour. Path, and Bact., 45, 1937, 75-78; Stuart -Harris, Brit. Jour. Exp. Path., 17, 1936, 324-328; 18, 1937, 485-492; Sulkin et al.. Jour. Inf. Dis., 69, 1941, 278-284; Tang, Brit. Jour. Exp. Path., 19, 1938, 179-183; Taylor, (A. R.), et al.. Jour. Immunol., Virus Res. and Exp. Chemother., 47, 1943, 261-282; Taylor, (R. M.), et al.. Am. Jour. Hyg., 31, (B), 1940, 36-45; Jour. Inf. Dis., 68, 1941, 90-96; Wells and Brown, Am. Jour. Hyg., 24, 1936, 407-413. 2. Tarpeia beta spec. nov. From sec- ond letter of Cireek alphabet. ('ommon name : Influenza B virus. Hosts : HOM I N I DAE^Homo sapiens L., man. Experimentally, also ferret, mouse, chick embryo. Geographical distribution: United States, England. Induced disease : In man, subclinical disease or one resembling that induced by influenza A virus. In chick embryo, ex- perimentally, virus increases in entoder- mal cells lining allantoic cavity. Serological relationships : Not neutral- ized by antiserum to influenza A virus. Specific neutralization and complement- fixation reactions. Rapidly adsorbed by normal chicken-blood red cells (95 per cent in 15 minutes) ; released in 4 hours essentially completely; the process is then repeatable with fresh red cells. Other properties : Particle circular or bean-shaped in outline, with average di- ameter of 97.3 millimicrons in electron micrographs ; of 99.8 millimicrons by centrifugation studies. Literature : Burnet, Austral. Jour. Exp. Biol, and Med. Sci., 19, 1941, 291-295; Francis, Science, 92, 1940, 405-408; Proc. Soc. Exp. Biol, and Med., 45, 1940, 861- 863; Hirst, Jour. Exp. Med., 76, 1942, 195-209; Lush et al., Brit. Jour. Exp. Path., 22, 1941, 302-304; Nigg et al.. Am. Jour. Hyg., 35, 1942, 265-284 ; Sharp et al.. Jour. Immunol., Virus Res. and Exp. Chemother., 48, 1944, 129-153. 3. Tarpeia premens spec. nov. From Latin premere, to oppress or afflict. Common name : Common-cold virus. Hosts : HOM I N I DAE— Homo sapiens FAMILY CHARONACEAE 127J L., man. Experimentally, also chimpan- zee, chick embryo. Geographical distribution : World-wide except in conditions of isolation of small communities. Induced disease : In man, incubation period about 48 hours ; mild malady ; run- ning nose in 81 per cent of cases, obstruc- tion of nostrils in 44 per cent, sudden onset in 37 per cent, cough in 31 per cent, headache in 19 per cent, sore throat in 14 per cent, fever in 13 per cent, inflamma- tion of eyes in 12 per cent ; changes in weather, especially during a warm season , predispose to the disease ; no correlation between susceptibility and outdoor exer- cise, exposure to fresh air while sleeping, eye color, adenotonsillectomy, or size of frontal sinus. Incidence inversely pro- portional to daily hours of sunshine and atmospheric temperature. Fitness (de- fined by speed of oxygen replacement ) correlated with relative freedom from colds. Effect of rest during disease fav- orable, reducing complications, length of fever, duration of illness, and period off duty. Immunological relationships: After at- tack, specific immunity for about 7 weeks (minimum period 23 days) ; then exposure to chilling maj^ cause a relapse, but an isolated community tends to lose the virus during the refractory period. Filterability : Passes Berkefeld V and W as well as Seitz filters. Other properties : Viable at least 13 days at ice-box temperature, anaerobi- cally ; at least 4 months frozen and dried in vacuo. Gum acacia tends to stabilize virus in chick-embryo tissue medium. Literature: Dochez et al., Jour. Exp. Med., 63, 1936, 559-579; Doull et al., Am. .Tour. Hyg., 13, 1931, 460-477; 17, 1933, 536-561; Gafafer, ibid., 13, 1931, 771-780; 16, 1932, 233-240, 880-884 ; Jour. Inf. Dis., 51, 1932, 489-492; Gafafer and Doull, Am. Jour. Hyg., 18, 1933, 712-726 ; Hyde and Chapman, ibid., 26, 1937, 116- 123; Kneeland et al., Froc. Soc. Exp. Biol, and Med., 35, 1936, 213-215; Le Blanc and Welborn, Am. Jour. Hyg., 2^, 1936, 19-24; Locke, Jour. Inf. Dis., 60, 1937, 106-112; Long and Doull, Proc. Soc. Exp. Biol, and Med., 28, 1930, 53-55; Maughan and Smiley, Am. Jour. Hyg., 9 1929, 466-472; Noble and Brainard, Jour. Bact., 29, 1935, 407-409; Palmer, Am. Jour. Hyg., 16, 1932, 224-232; Paul and Freese, ibid., 17, 1933, 517-535; Shibley et al.. Jour. Am. Med. Assoc, 95, 1930, 1553-1556; Smiley, Am. Jour. Hyg., 6, 1926, 621-626; 5, 1929, 477-479. 4. Tarpeia fells .s'pec. noiJ. From Latin feles, cat. Common name : Feline-distemper virus. Hosts : FELIDAE—Felis catus L., do- mestic cat ; F. pardus, leopard ; F. ligrina, American tiger cat ; F. aurata, African tiger cat; F. planiceps, rusty tiger cat; F. marmorata, marbled cat ; F. caracal, caracal lynx; F. pardalis, ocelot; lion, tiger, puma relatively insusceptible. Insusceptible species : Man, dog, ferret , mongoose, rabbit, rat, mouse, guinea pig. Induced disease: In domestic cat, coughing, sneezing, running eyes and nose, with serous or purulent conjunctivi- tis, or diarrhea and vomiting; fever to 103 or 105° F; loss of appetite, general weakness; mortality high, especially among j'oung individuals; death usually occurs on the 10th to the 12th day, in extreme cases, however, as early as the 5th or as late as the 35th day; catarrhal congestion in some part of the gastro- intestinal tract is typical; this ranges from a fev: small patches in the ileum to involvement of the whole small intes- tine and parts of the large intestine or stomach and esophagus; often enlarge- ment and congestion of abdominal lymph glands, enlargement of spleen, pleurisy, and peritonitis. Filterability : Passes Berkefeld N and Chamberland L3 filters. Transmission : By fomite s. Imnmnological relationships : Recov- ered cats specifically immune. 1272 MANUAL OF DETERMINATIVE BACTERIOLOGY Other properties : Viable at least 3 weeks in 50 per cent glycerine ; attenu- ated or killed by drying at room tempera- ture, but some immunization is reported if dried virus is injected. Literature: Dalling, Vet. Record, 15, 1935, 283-289; Findlay, Vet. Jour., 89, 1933, 17-20; Hindle and Findlay, Jour. Comp. Path, and Therap., 45, 1932, 11-26 ; Verge and Cristoforoni, Compt. rend. Soc. Biol., Paris, 99, 1928, 312-314. 5. Tarpeia vitulae spec. Jtuv. From Latin vitula, cow-calf. Common name : Pneumoenteritis virus Hosts : BOVIDAE—Bos taurus L., do- mestic cattle. Experimentally, also MURIDAE — Mus musculus L., mouse. Geographical distribution: United States. Induced disease: In cattle (calves), after incubation pei'iod of 2 to 4 days, fever increasing rapidly to 40 or 41° C and lasting 3 to 5 days ; usually after first day of fever, diarrhea with feces soft, yellow, voluminous, fetid in odor, occa- sionally blood-tinged or fluid; diarrhea is followed by pneumonia and recovery after disappearance of fever; catarrhal enteritis and a bronchopneumonia usually confined to the anterior lobes of the lungs underlie the symptoms; no inclusion bodies in cells of affected tissues. Transmission: By pen contacts with infected calves. E.xperimentally, by in- tranasal inoculation of calves, using in- ocula prepared from lungs of infected mice. Serological relationships : Recovered animals develop neutralizing antibodies. Immunological relationships : A specific resistance to reinfection is conferred by an attack of the disease. Filterability : Passes Berkefeld N filter. Literature: Baker, Cornell Vet.,' 32, 1942, 202-204; Jour. Exp. Med. 78, 1943, 435-446. 6. Tarpeia canis spec. nov. From La- tin canis, dog. Common name : Canine-distemper virus. Hosts: CAN I DAE — Canis familiaris L., dog; Viilpes sp., fox. MUSTELI- DAE—iQi-vei. Insusceptible species: Man, rabbit, guinea pig, white rat, cat. Geographical distribution : Widespread throughout the world. Induced disease : In dog, after 4 days from time of infection, fever and a watery discharge from the eyes and nose, some- times inconspicuous but often profuse ; usually diarrhea and wasting followed by recovery or, exceptionally, death. Virus passes from the respiratory tract through the blood stream to its favored sites in vascular endothelium and cells of the reticulo-endothelial system. Nuclear in- clusions are found in liver cells, bronchial epithelial cells, glandular cells of the stom- ach and intestine, and bile-duct epithe- lial cells; there are also cytoplasmic inclusions in bile-duct epithelial cells. Transmission: By contact. Probably by air-borne droplets. No arthropod vector is recognized. Immunological relationships : Dead- vaccine treatment followed by living- virus treatment produces a lasting im- munity. Virus inactivated by photo- dynamic effect in 2 mm layer of 1 :50,000 or 1 : 100,000 methylene blue, exposed 30 minutes at 20 cm from 100 candle-power lamp, still immunizes. Vaccine may be dried. Filterability: Passes Chamberland L2 and Mandler filters. Other properties : Viable in liver tissue at 10° C for 35, not 85, days ; in glycerine- saline solution at 10° C, 67 days though deteriorated; in vacuum-dried liver tis- sue, at 10° C, 90 days. If dried from frozen state, virus is viable in vacuum at least 430 days at 7° C, in oxygen-free nitrogen at least 365 days at 7° C. Viable in 25 per cent sterile horse serum at —24° C more than 693 days. Literature : Carre, Compt. rend. Acad. Sci., Paris, HO, 1905, 689-690; Dalldorf, FAMILY CHARONACEAE 1273 Jour. Exp. Med., 70, 1939, 19-27; De Monbreun, Am. Jour. Path., 13, 1937, 187-212; Dunkin and Laidlaw, Jour. Comp. Path, and Therap., 39, 1926, 201- 212. 213-221; Green and Evans, Am. Jour. Hyg., 29 (B), 1939, 73-87; Laidlaw and Dunkin, Jour. Comp. Path, and Therap., 39, 1926, 222-230; 41, 1928, 1-17, 209-227; Perdrau and Todd, ibid., 46. 1933, 78-89; Siedentopf and Green, Jour. Inf. Dis., 71, 1942, 253-259 ; Wharton and Wharton, Am. Jour. Hyg., 19, 1934, 189-216. 7." Tarpeia vulpis .spec. nov. From Latin vidpes, fox. Common name : Fox-encephalitis virus. Hosts : CAN I DAE — Vvlpes sp., silver fox. Experimentally, also some, but not all, dogs; coyote. Insusceptible species : Gray fox, mink, ferret, sheep, laboratory rabbit. Geographical distribution: Unite I States. Induced disease: In fox. after 2 daj's from time of infection, loss of appetite, slight nasal discharge ; convulsions with earl}^ death or h.yperexcitabilitj', blind walking, lethargy, flaccid or spastic paral- ysis, muscular twitching, tearfulness, weakness, coma and death; many more foxes become infected in epizootics than show obvious disease, some being symp- tomless carriers ; 12 to 20 per cent fatali- ties may be experienced among j'^oung foxes on ranches, 3 to 9 per cent among adults. Intranuclear inclusions in vascu- lar endothelial cells especially in cerebral endothelium; sometimes in hepatic cells and endothelial cells of liver and kidney ; no intracytoplasmic inclusions; virus in heart blood, spleen, and brain ; in carriers, virus is believed to persist in focal lesions in upper respiratory tract. Experimen- tallj' in susceptible dogs, sometimes coryza, discharge from eyes and nose often purulent, commonly fits of excite- ment, coma, death; recovery rare; cellu- lar infiltration in the central nervous sj^s- tem, focal necrosis of the liver; specific intranuclear inclusions in cells of the vascular endothelium, meningeal cells, reticulo-endothelium, hepatic cells, and occasionally in cortical cells of the adrenal. Transmission : Experimentally, by skin scarification, intramuscular injection, in- traperitoneal injection, inoculation of cisterna. intratesticular injection, inocu- lation of nasal cavity; not by corneal scarification. Immunological relationships : Injec- tions of this virus afford no immunity to subsequent infection by canine distemper virus. Filterabilit J' : Passes Berkefeld X filter. Other properties : Viable in 50 per cent glycerine for several j'ears, in carcass for several days. Literature: Barton and Green, Am. Jour. Hyg., 37, 1943, 21-36; Green, Proc. Soc. Exp. Biol, and Med., 23, 1926, 677- 678; Am. Jour. Hyg., 13, 1931, 201-223; Green and Dewey, Proc. Soc. Exp. Biol, and Med., 27, 1929, 129-130; Green and Evans, Am. Jour. Hyg., 29 (B), 1939, 73-87; Green and Shillinger, ibid., 19, 1934, 362-391 ; Green et al., ibid., 12, 1930, 109-129; 14, 1931, 35.3-373; 18, 1933, 462-481 ; 19, 1934, 343-361 ; 21, 1935, 366- 388; 24, 1936, 57-70; Lucas, Am. Jour. Path., 16, 1940, 7.39-760. 8. Tarpeia viverrae spec. nor. From Latin viverra, ferret. Common name: Ferret -distemper virus. Host : MUSTELIDAE—Mnstelafuro. ferret. Insusceptible species : Dog, mouse, rat, guinea pig, rabbit. Geographical distribution: United States. Induced disease: In ferret, fever to 105 or 106° F, lethargy, loss of appetite, conjunctivitis with exudate closing eyes, sometimes a purulant nasal discharge, weight loss small, sneezing rare, difficulty in breathing, death 14 to 56 days after inoculation (average 20 days), sometimes 1274 MANUAL OF DETERMINATIVE BACTEfilOLOGf preceded by convulsions and other nerv- ous signs ; fatality rate 70 to 100 per cent. Transmission: By cage contacts. By feeding. Experimentally by intranasal, subcutaneous, or intradermal inoculation. Immunological relationships: In im- munized animals, no cross immunity with canine distemper virus nor with human influenza virus. Thermal inartivation : At 60° C in 30 minutes. Filterability : Passes Berkefeld N filter. Other properties : Viable at least 3, but not 5, months in 50 per cent neutral gly- cerine ; at least 4 months when frozen and dried in vacuo. Literature : Slanetz and Smetana, Jour. Exp. Med., 66, 1937, 653-666; Spooner, Jour. H3^g., 38, 1938, 79-89. 9. Tarpeia avium s-pec. nov. From Latin aves, fowl of tlie air. Common names : Laryngotracheitis virus; also known as infectious laryn- gotracheitis virus and as infectious bron- chitis virus. Hosts : PHASIANIDAE—Gallus gal- lus (L.), chicken. Experimentally, also PHASIAN ID AE— pheasant ; Fi hybrid between male Ringneck pheasant and female bantam chicken ; chorioallantoic membrane of developing chicken embryo (with macroscopic lesions on membrane as a result of proliferative and necrotic changes) ; turkey embryo. Insusceptible species : Guinea fowl (no evidence of disease on inoculation) ; white rat, guinea pig, rabbit ; embr\'Os of pigeon, guinea fowl, and duck. Geographical distribution : United States, Canada, Australia. Induced disease : In domestic chicken, mostly among pullets and yearling hens, loss of appetite, lachrymation from one or both eyes, respiratory distress, hemor- rhagic and mucous exudate in lumen of trachea and occasionally^ in t he bronchi ; death as a result of asphyxiation or, more often, recovery; recovered birds occa- sionally carry the virus in the upper respiratory tract for some time (a period of 467 days has been recorded) ; virus is not found on eggs during an outbreak in a flock, but is always in trachea of an affected bird; intranuclear inclusions in tracheal lesions ; virus has special affinity for mucous membrane of eye, nostril, larynx, trachea, cloaca, and bursa of Fabricius ; usually affects more than half the birds in a flock, with a mortality of 5 to 60 per cent (averaging between 10 and 20 per cent). Transmission : By contacts. Experi- mentally, by intrabursal injection (in bursa of Fabricius) or by rubbing the mucous membrane in the dorsal region of the outer or proctodeal part of the cloaca with a small cotton swab moistened with a suspension of virus. Serological relationships : Serum from recovered fowl neutralizes virus ; dilution tends to reactivate neutralized virus. Immunological relationships : Experi- mental infection of cloaca and bursa of Fabricius, especially in 2 to 4-month-old birds, immunizes against infection by subsequent tracheal inoculation. Thermal inactivation : At 55.5° C in 10 to 15 minutes ; at 60° C in 2 to 3 minutes ; at 75° C in J to i minute ; all tests with virus in the presence of tracheal exudate. Filterability : Passes Berkefeld V and N filters. Strains : A Victorian strain has been re- ported as of low virulence for fowls. Other properties : Inactivated in 5 per cent phenol in 1 minute ; in 3 per cent cresol compound in | minute ; in 1 per cent sodium hydroxide in ^ minute. Viable in tracheal fluid in dark for 75, not 110, days ; in light for 6, not 7, hours ; in buffer solu- tion at pH 7.4 for 131 days ; at 4 to 10° C in dark for at least 217 days ; in dried state for at least 661 days. Viable in dead body at 37° C for 22, not 44, hours ; at 13 to 23° C for 10, not 15, days; at 4 to 10° C for 30, not 60, days. Literature : Beach, Science, 72, 1930, 633-634; Jour. Exp. Med., 54, 1931, 809- 816; Jour. Inf. Dis., 57, 1935, 133-135; FAMILY CHARONACEAE 1275 Beach et al., Poultry Science, IS, 1934, 218-226; Beauclette and Hudson, Science, 76, 1932, 34; Jour. Am. Vet. Med. Assoc, 82 (X.S. 35), 1933, 460-476; 55, 1939, 333- 339; Brandly, ibid., 88 (N.S. 41), 1936, 587-599; Jour. Inf. Dis., 57, 1935, 201- 206; Braudly and Bushnell, Poultry Science, 13, 1934, 212-217; Burnet, Brit. Jour. Exp. Path., 15, 1934, 52-55; Jour. Exp. Med., 63, 1936, 685-701 ; Burnet and Foley, Austral. Jour. Exp. Biol, and Med. Sci., 19, 1941, 235-240; Gibbs, Jour. Am. Vet. Med. Assoc, 81, (N.S. 34), 1932, 651-654; Massachusetts Agr. Exp. Sta., Bull. 295, 1933, ibid., Bull. 311, 1934; Hinshaw et al., Poultry Science, 10, 1931, 375-382; Hudson and Beaudette, Science, 76, 1932, 34; Cornell Vet., 22, 1932, 70-74; Kernohan, California Agr. Exp. Sta., Bull. 494, 1930, 3-22 ; Jour. Am. Vet. Med. Assoc, 78 (N.S. 31), 1931, 553-555; Komarov and Beaudette, Poultry Sci- ence, 11, 1932, 335-338 ; May and Tittsler, Jour. Am. Vet. Med. Assoc, 67, (N.S. 20), 1925, 229-231; Schalm and Beach, Jour. Inf. Dis., 56, 1935, 210-223; Seifried, Jour. Exp. Med., 54, 1031, 817-826. Gentis III. Tortor gen. nov. Viruses of the Hog-Cholera Group, inducing diseases characterized by involvemer of manj'^ tissues. Generic name from Latin tortor, tormentor. The type species is Tortor suis spec. nov. Key to the species of genus Tortor. I. In roammals. A. Infecting swine. B. Infecting cattle. C. Infecting the horse. D. Infecting sheep. E. Infecting cat. II. In birds. 1. Tortor suis spec. nov. From Latin sus, hog. Common names : Hog-cholera virus, swine-fever virus. Host: SU I DAE— Sus scrofa L., do- mestic swine. Wart hog (symptomless carrier). Insusceptible species: Dog, cat, cow, horse, donkey, sheep, goat, rabbit, guinea pig, mouse, rat, goose, hen, duck, pigeon. Geographical distribution : Almost uni- versal in pig-breeding countries, espe- 1. Tortor suis. 2. Tortor bovis. 3. Tortor equorvm. 4. Tortor equcs. 5. Tortor ovis. 6. Tortor felis. 7. Tortor galli. 8. Tortor fur ens. cially in Europe, the British Isles, North and South America. Induced disease : In swine, after intra- muscular injection, increased tempera- ture and prostration within 2| to 3 days ; later lymph nodes enlarged, sometimes hemorrhagic ; hemorrhages under capsule of kidneys. Virus may remain in blood of recovered pigs for 10 months. Ac- quired immuuit}' is lasting, but most naturally infected animals die in newly infected herds. Virus has been cultured 1276 MANUAL OF DETERMINATIVE BACTERIOLOGY in minced swine testicle on solid serum- agar and on egg membrane, increase being limited to the living tissues from the swine and furnishing inoculum active in amounts as small as 10~^ ml. Transmission : By feeding. Through air contamination. Rarely by contact. Experimentally, by subcutaneous injec- tion. Urine highly infective. Virus in blood and all tissues early in disease. Serological relationships : Immune serum affords passive protection. Thermal inactivation : At 55° C in 30 minutes ; at 60° C in 10 minutes. At 72° C in 1 hour in dried blood. Filterability : Passes Berkefeld filter. Other properties : Viable in blood in cool, dark place at least 6 years. Literature : De Kock et al., Onder- stepoort Jour. Vet. Sci. and Anim. In- dust., 14-, 1940, 31-93; Hecke, Cent. f. Bakt., I Abt., Orig., 126, 1932, 517-526; Montgomery, Jour. Comp. Path, and Therap., Sit, 1921, 159-191 ; Rohrer, Arch. Tierheilk., 62, 1930, 345-372, 439-462; 6J^, 1931, 124-143; TenBroeck, Jour. Exp. Med., 74, 1941, 427-432. 2. Tortor bovis spec. nov. From Latin bos, cow. Common names : Cattle-plague virus, virus of pestis bovina, runderpest virus, Rinderpest virus. Hosts : BOVIDAE—Bos tavrus L., do- mestic cattle ; swine, buffalo, zebu cattle, sheep, goat , camel, deer. Koedoe, eland, bushbuck, duiker, and other antelopes. Insusceptible species: Man, solipeds, carnivora. Geographical distribution : Widespread over Asia and the Asiatic islands. At times in Western Europe. Enzootically in Turkey. Periodically in North Africa, especially in Egypt ; at times throughout Africa. Not in North America. At times in South America, Australia (sup- pressed quickly). Induced disease: In domestic cattle, after 3 to 9 days, febrile reaction, restless- ness, loss of appetite, cessation of rumina- tion ; fever highest at 5th or 6th day of disease, then temperature drops to normal or subnormal and diarrhea begins ; muzzle dry, coat staring, hair dull, skin moist in parts; twitching of superficial muscles, grinding of teeth, arching of back, glairy discharge from nose, redness of mucous membranes ; restlessness increases, diar- rhea becomes severe with fetid, blood- stained or blackish liquid discharges ; weakness, drooping of ears, occasional yawning, coldness of extremities; occa- sionally excitement precedes weakness ; skin may become red and moist, showing protuberances and vesicles, with matted hair ; later wrinkling and scab formation ; conjunctiva red, eyelids swollen, tears flowing, followed by mucous, then puru- lent, discharge; sometimes a cough develops and respirations become rapid; red spots inside mouth develop into ero- sions or ulcers, often confluent ; pregnant animals often abort; milk of cows de- creases, sometimes becoming yellow and watery. Death is sometimes early (1 to 2 days after first manifestations of dis- ease), more often delayed (4 to 7 days) ; .sometimes animals live 2 or 3 weeks or longer. Disease milder and more chronic where enzootic ; morbidity to 100 per cent and mortality to 96 per cent in new areas. Recovered animals show a lasting, sterile mmunity. Urine, feces, nasal and lach- rymal discharges, sweat, aqueous humour, cerebrospinal fluid, lymph, emulsions of viscera and muscles, and blood are infec- tive during the course of the disease. Transmission : By contact, even during prodromal period; by contaminated food, troughs, or other articles. No insect vector is known. Immunological relationships : One at- tack confers a lasting immunity, except rarely, when a mild second attack may occur. A calf from a diseased mother may be resistant if pregnancy was far advanced when the disease occurred. Filterability : Passes Berkefeld V filter candle, with difficult J^ Other properties : Remains infective at FAMILY CHARONACEAE 1277 least 2 weeks at 0° C in virulent blood, less than 2 days in hides dried in direct sunlight, 3 days in contaminated wool, as long as 12 days in meat ; is inactivated by glycerine, bile, chloroform, formalin, and 2 per cent phenol ; is virulent at least 25 days in body of leech, Hirudo boynioni Wharton (HIRUDIDAE), fed on sick animal. Literature : Boynton, Philippine Agr. Rev., 10, 1917, 410-433; Daubney, Jour. Comp. Path, and Therap., 41, 1928, 228- 248; 263-297; Hornby, ibid., 41, 1928, 17- 24; Pfaff, Onderstepoort Jour. Vet. Sci. and Anim. Indust., 11, 1938, 263-330; 15, 1940, 175-184; Weston, Jour. Am. Vet. Med. Assoc, 66 (N.S. 19), 1924, 3.37-,350. 3. Tortor equorum spec. nov. From Latin eqinis, horse. Common names : Horse -sickness virus, African horse-sickness virus, virus of pestis equorum, virus of perdesiekte, virus of South African Pferdesterbe. Hosts: EQUIDAE—Equus caballus L., horse; perhaps E. asinus L., donkey. Experimentally, also CA N IDAE — Canis familiaris L., dog. CA VI IDAE — Cavia ■porcellus (L.), guinea pig. MURIDAE — Rattus norwegicus (Erxleben) , wild and albino rat ; mouse ; Angora goat ; Mastomys concha, multimammate mouse ; Tatera lobengula, gerbille ; chick embryo (but no virus in hatched chick). Mule and zebra relatively resistant. Insusceptible species: HOMI NIDAE — Homo sapiens L., man. LEPORI- DAE- — Oryctolagus cunicnhis (L.), rabbit (no observed disease). Geographical distribution : Africa, es- pecially in coastal regions and river valleys. Induced disease : In the horse, four types of disease are recognized. Horse- sickness fever, prodromal period 5 to 28 days, rise of body temperature to 105° F in 1 to 3 days, with return to normal temperatures in another day or two; sometimes loss of appetite, redness of conjunctiva, labored breathing, and ac- celerated pulse ; recovery prompt . Dun- kop or acute pulmonary horse -sickness, prodromal period of 3 to 5 days, severe dyspnea, fever, coughing, frothing at nostrils; fever to 106° F, breathing rate to 60 a minute, nostrils dilated, head and neck extended, ears drooping, sweating, progressive weakness; often fatal. Dik- kop, or cardiac form of horse-sickness, jjrodromal period 5 to 21 days, fever develops slowly, lasts long; edematous swellings of head and neck, symptoms of cardiac dyspnea, sometimes blood spots on conjunctiva, mucous membranes of mouth and tongue bluish, restlessness ; sometimes fatal outcome. Mixed form of horse-sickness, combining features of pulmonary and cardiac types. Horses recovering from natural infections are known as "salted" and possess heightened resistance to the disease. Transmission: Not by contact. Mos- quitoes and biting flies have been sus- pected as vectors. Experimentally, bj^ intravenous or subcutaneous injection. Serological relationships : Serologically distinguishable strains exist. Immunological relationships : Immun- ity to liomologous strain complete after an attack (horse then known as "salted" for that strain), but immunity to hetero- logous strains incomplete. Antibodies absent from young at birth but as high in titer as in dam within 30 hours, presum- ably from colostral milk ; declining gradu- ally over a period of about 6 months. Thermal inactivation : At 57.5 to 60° C in 10 minutes. Filterability : Passes Berkefeld, Cham- berland F, and Seitz EK filters. Other properties : Viable dry at least 15 months. Stable in alkaline solutions (to pH 10), unstable in acid (beyond pH 6.0). Serum-saline solutions preferable to saline solutions for storage. Particle diameter determined as 40 to 60 milli- microns (mean 50 millimicrons) by filtra- tion methods, 45.4 millimicrons by cen- trifuging. Density 1.25 gm per ml. Isoelectric point at pH 4.8. 1278 MANUAL OF DETERMINATIVE BACTERIOLOGY Literature : Alexander, Onderstepoort Jour. Vet. Sci. and Anim. Indust., 4, 1935, 291-322, 323-348, 349-377, 379-388; 7, 1936, 11-16; U, 1938, 9-19; Alexander and DuToit, ibid., 2, 1934, 375-391; Alexander and Mason, ibid., 16, 1941, 19-32; Alexander et al., ibid., 7, 1936, 17-30; DuToit et al., ibid., 1, 1933, 21-24, 25-50; Henning, in Animal Diseases in South Africa, Central News Agencj', Limited, South Africa, 2, 1932, 516-538; M'Fadyean, Jour. Comp. Path, and Therap., 13, 1900, 1-20; 23, 1910, 27-33, 325-328; Nieschulz and DuToit, Onder- stepoort Jour. Vet. Med. and Anim. Indust., 8, 1937, 213-268; Poison, ibid., 16, 1941, 33-50, 51-66; Nature, H8, 1941, 593-594; Theiler, Deutsch. tierarztl. Wochenschr., 9, 1901, 201-203, 221-226, 233-237, 241-242 ; Report for 1905-1906 of the Govt. Veterinary Bacteriologist, Transvaal Dept. Agr., 1907, 160-162; Jour. Comp. Path, and Therap., 23, 1910, 315-325. 4. Tortorequaes73ec.no;'. From Latin equa, mare. Common name : Mare-abortion virus. Hosts: EQUIDAE — Equus caballus L., horse. Experimentally, also Syrian hamster (newborn); tissues of human placenta grafted on the chorioallantois of the chick embryo. Insusceptible species : Chicken (em- bryo ; no observed susceptibility). Induced disease : In horse, small, mul- tiple, grayish white areas of necrosis in the livers of aborted fetuses; acidophilic intranuclear inclusions in hepatic cells around these foci, in epithelial cells of bile ducts, and in bronchial epithelium; petechial hemorrhages in the heart, spleen, and lungs; excess fluid in the thoracic cavity. Transmission: By contact. By living in contaminated stalls. Literature : Anderson and (ioodpasture, Am. Jour. Path., 18, 1942, 555-561; Dimock, Jour. Am. Vet. Med. Assoc, 96, 1940, 665-666; Dimock and Edwards, Cornell Vet., 26, 1936, 231-240; Goodpas- ture and Anderson, Am. Jour. Path., 18, 1942, 563-575; Hupbauer, Munch. Tier- arztl. Wchnschr., 89, 1938, 37-38; Miess- ner and Harms, Deutsche Tierarztl. Wchnschr., 46, 1938, 745-748. 5. Tortor ovis spec. nov. From Latin ovis, sheep. Common name : Blue-tongue virus. Hosts: BOVIDAE—Ovis aries L., sheep; Bos taurus L., cattle. Geographical distribution : South Africa. Induced disease : Both sheep and cattle may carry the virus at times without obvious manifestations of disease or there may be severe manifestations. In sheep, experimentally, diffuse hyperemia of buccal mucosa, especially of lips ; then petechiae and ecchymoses followed by excoriations and necrosis of the mucous membrane, especially on lips, tongue, inside of cheeks, dental pad, gums, muz- zle, and external nares; sometimes deep seated necrotic ulcers on tongue de- veloping from the more usual superficial necrotic process ; mucoid discharge from nostrils, becoming muco-hemorrhagic ; commonly frothing at the mouth in early stages of the disease; frequently redden- ing of skin of lips and nose ; rarely whole skin becomes flushed and wool is shed; often swelling of vulva with necrotic changes on borders and petechiae in mucosa ; tongue sometimes swollen ; lame- ness common and severe; recovery or death. In cattle, edema of lips and tongue; hyperemia of oral mucosa; mul- tiple hemorrhages in skin, lips, mucous membrane of the lips, tongue, dental pad, buccal cavity, small intestine, myocar- dium, epicardium, and endocardium, less frequently in the trachea, nasal cavity, bladder, urethra, pulmonary artery, and pleura; localized necrotic areas followed by ulceration on lips, gums, the dental pad, tongue, mucous membrane of the rumen, pylorus of the stomach, and the external nares ; scattered skin lesions with reddening, slight exudation, crusting, sloughing of crusts and hair together, FAMILY CHARONACEAE 1279 mucoid or mucopurulent discharge from nostrils; prognosis favorable in mild cases, but disease occasionally terminates with death. Transmission: Not by contact; arth- ropod vector suspected. Other properties : Infective particle calculated to be 87 to 105 millimicrons in diameter by sedimentation studies, 100 to 132 millimicrons in diameter by ultra- filtration. Literature: Bekker et al., Onderste- poort Jour. Vet. Sci. and Anim. Indust., 2, 1934, 393-507; De Kock et al., ibid., 8, 1937, 129-180; Henning, in Henning, M. W., Animal Diseases in South Africa, Central News Agency, Ltd., South Africa, 1932, vol. 2, chapter 27, pages 503-515 ; Mason and Neitz, Onderstepoort Jour. Vet. Sci. and Anim. Indust., 16, 1940, 149-157; Nieschulz et al., ibid., 2, 1934, 509-562; Poison, Nature, ^^8, 1941, 593-594. 6. Tortor felis spec. nov. From Latin feles, cat. Common names : Panleucopenia virus, infectious feline agranulocytosis virus, in- fectious aleucocytosis virus, feline en- teritis virus. Host: FELIDAE— Felis calus L., do- mestic cat. Insusceptible species : White mouse, guinea pig, domestic rabbit, ferret; Citellus richardsonii (Sabine), ground squirrel. Geographical distribution : United States, Germany. Induced disease : In cat, variable ef- fects, some individuals little affected, others listless, recumbent, refusing food, showing some vomiting, diarrhea, nasal and ocular discharges; often death, after a few minutes of fibrillary twitching and terminal clonic convulsions, before there is much loss of weight; sometimes recov- ery with return of appetite. Profound leucopenia and marked relative lympho- cytosis without thrombopenia or appreci- able anemia; proliferation of reticulo- endothelial cells of lymph nodes and spleen ; intranuclear inclusion in cells of gastrointestinal mucosa, spleen, lymph nodes, bone marrow, and bronchial mucosa. Transmission : Perhaps by nasal drop- lets or contaminated food. No arthropod vector recognized. Experimentally by oral, intragastric, cutaneous, subcutane- ous, intraperitoneal, intravenous, and intranasal routes. Serological relationships : Sera from panleucopenia-immune cats protects against agranulocytosis virus. Immunological relationships : Cats im- mune as a result of earlier infection with agranulocytosis virus resist later inocula- tion with panleucopenia virus. Previous inoculation ineffective if made with hog- cholera virus or fox-encephalitis virus. Filterability : Passes Berkefeld V, N, and W filters and Seitz EK discs. Other properties : Remains active in 50 per cent glycerine at least 138 days in tissues ; not inactivated by drying while frozen, nor by freezing at about —80° C. Literature : Hammon and Enders, Jour. Exp. Med., 69, 1939, 327-352; 70, 1939, 557-564; Kikuth et al., Cent. f. Bakt., I Abt., Orig., 146, 1940, 1-17; Lawrence and Syverton, Proc. Soc. Exp. Biol, and Med., 88, 1938, 914-918; Lawrence et al.. Jour. Exp. Med., 77, 1943, 57-64; Am. Jour. Path., 16, 1940, 333-354; Syverton et al.. Jour. Exp. Med., 77, 1943, 41-56. 7. Tortor galli spec. nov. From Latin gallus, cock. Common names : Fowl-plague virus, fowl-pest virus. Hosts: Chiefly chicken, turkey, goose. Experimentally, also ferret, rhesus mon- key, hedgehog, pigeon, duck, canary, mouse, rat, rabbit. Multiplies in em- bryonated hen's egg; edema, but no dis- crete primarj^ lesions in chorioallantoic membrane. Geographical distribution : Widespread throughout Europe. North and South America, Asia. Induced disease : In chicken, loss of appetite, tendency to leave companions 1280 MANUAL OF DETERMINATIVE BACTERIOLOGY and seek shade, drooping of wings and tail ; eyes closed or partly closed ; some dyspnea; in some cases, edema of head and neck; in late stages, sometimes cyanosis of comb and skin; staggering, twitching, or spasms; fever may disap- pear and temperature become subnormal before death; recovery in about 30 per cent of all cases ; linear and punctiform hemorrhages throughout body. Transmission : Method of natural trans- mission unknown. The fowl louse, Goni- oides dissimilis {PHILOPTERIDAE), has been suspected as vector (Maggiora and Tombolato, Rendiconti, Accademia delle Scienze dell'Instituto di Bologna, n.s. 27, 1923, 200-203). Experimentally, by subcutaneous, intramuscular, and intravenous injection. Serological relationships : Specific neu- tralizing antiserum does not react with influenza virus. No reaction of fowl- plague virus with antisera specific for canine distemper, influenza, or Rift Valley fever viruses. Thermal inactivatioii : At 55° C in 1 hour in whole blood or brain. Filterability : Passes membrane of average pore diameter 150, not 100, not ordinarily 125, millimicrons. Passes Berkefeld and Chamberland filters. Other properties : Particle diameter es- timated by filtration as 60 to 90 millimi- crons; by centrifugation, as 120 to 130 millimicrons. Viable after exposure in 1 : 10,000 dilution for 10 minutes, in 2 mm layer of 1:50,000 methylene blue, 15 cm from a 300 candle-power filament lamp. Withstands drjdng. Precipitates from salt-free solutions or in presence of half- saturated ammonium sulphate solutions; virus held to be of globulin nature by Mrowka, Cent. f. Bakt., I Abt., Ov\g., 67, 1912, 249-268. Strains : Variant strains have been produced by intracerebral passage in brains of canaries and mice. Literature : Bechhold and Schlesingcr, Biochem. Ztschr., 236, 1931, 387-414; Ztschr. Hyg. Infektionskr., 112, 1931, 668-679; Burnet and Ferry, Brit. Jour. Exp. Path., 15, 1934, 56-64; Centanni, Cent. f. Bakt., I Abt., Orig., 31, 1902, 145-152, 182-201 ; Elford and Todd, Brit. Jour. Exp. Path., U, 1933, 240-246; Findlay and Mackenzie, ibid., 18, 1937, 146-155, 258-264; Findlay et al., Jour. Path, and Bact., 45, 1937, 589-596 ; Lepine Compt. rend. Soc. Biol., Paris, 121, 1936, 509-510; Mackenzie and Findlay, Brit. Jour. Exp. Path., 18, 1937, 138-145; Nieschulz and Bos, Cent, f . Bakt., I Abt., Orig., 131, 1934, 1-6; Plotz and Haber, Compt. rend. Soc. Biol., Paris, 125, 1937, 339-340. 8. Tortor furens spec. nov. From Latin /wrere, to rage. Common name : Newcastle -disease virus. Hosts : PHASIANIDAE—Gallus gal- lus (L.), domestic chicken. HOMINI- DAE — Homo sapiens L., man (by labora- tory accident). Experimentally, also pigeon; chick embryo (with primary lesions and cytoplasmic inclusions in chorioallantoic membrane ) . Geographical distribution : England, probably also East Indies, Korea, Japan, Lidia, Australia. Induced disease : In chicken, acute, febrile, highly contagious, usually fatal disease resembling fowl plague ; loss of appetite, crouching attitude, half closed eyes, rapid respirations, waterj^ yellow- ish-white diarrhea with nauseating odor ; death usuallj' between 6th and 8th day. In man, accidentally infected in labora- tory by virus sprayed into eye, virus recoverable from temporarily inflamed eye ; recovery in 8 days with gradual in- crease of specific antibodies in blood. Transmission : By contact between liealthy and diseased birds. Serological relationships : Antiserum effective in neutralizing homologous virus. Immunological relationships : Chickens immune to infection by fowl-plague virus are susceptible to infection by this virus FAMILY CHARONACEAE 12SI and vice versa. Immunization to this virus does not decrease susceptibility to comb or mouth form of fowl pox. Thermal inactivation : At 60° C in 1 hour ; not at 56° C in 30 minutes. Filterability : Passes Berkefeld, Cham- berland L3, and Seitz filters. Other properties : Particle diameter calculated from filtration experiments to be 80 to 120 millimicrons. Not inacti- vated in 30 minutes in 1 ; 50,000 methylene blue solution in 2 mm layer 15 cm from a 300 candle-power filament lamp. Literature : Burnet, Med. Jour. Aus- tralia, 30, 1943, 313-314; Burnet and Ferry, Brit. Jour. Exp. Path., 15, 1934, 56-64; Doyle, Jour. Comp. Path, and Therap., 40, 1927, 144-169. 1282 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY V. TRIFURACEAE FAM. NOV. Viruses of the Infectious Aneniia Group, inducing diseases niainly characterized by- disturbances in balance of blood cells. There is a single genus. Genus Trifur gen. nov. With characters of the family. Generic name from Latin trifur, arrant thief. The type species is Trifur equorum spec. nov. I. Affecting horse. II. Affecting fowl. Key to the species of genus Trifur. 1. Trifur equorum. 2. Trifur gallinarum^ 1. Trifur equorum spec. nov. From Latin equus, horse. Common name : Equine infectious- anemia virus. Hosts: EQU I DAE— Equus caballus L., horse; E. asinus L., donkey. HO- MINIDAE — Homo sapiens L., man. Experimentally, also EQU I DAE— Equus asinus X E. caballus, mule. SUIDAE — Sus scrofa L., swine. Insusceptible species: BOVIDAE — Bos taurus L., cattle; Ovis aries L., sheep; Capra hircus L., goat. CANI- DAE — Canis familiaris L., dog. Geographical distribution : Europe, Union of South Africa, United States, Canada, Japan; at times in most parts of the world; not Australia. Induced disease : In horse, progressive anemia with eventual death or clinical recovery and retention of virus; disease may be acute, subacute, or chronic; in acute disease, temperature rise to 104 to 105° F. or even 106 to 107° F, remaining high much of the time until death or change to subacute or chronic form; in the acute form of the disease there is dull- ness, decreased appetite, drooping of head, flexing of limb not supporting weight; sometimes increase in pulse frequency to 70 or even 100 a minute but oftener rates around 50 a minute ; con- junctiva sometimes colored orange, with injection of vessels and petechiae, later becoming muddy colored or pale red, membrane, edematous; uncertain gait, trailing of hind feet, prostration, some- times death; subacute disease milder and with remissions ; chronic disease stilt milder, anemia conspicuous, sometimes death from debility or at end of a febrile attack; blood infective long (3 to 7 years) after clinical recovery; urine infective to horse by mouth. In man, diarrhea al- ternating with constipation, herpes-like exanthema on abdominal wall, blood sometimes in feces ; persistent headache, temperature normal ; later, lumbar pains, generalized edema, general debility, loss of flesh, pallor of face and mucosae; fil- tered blood in 1 ml. amount fatal to horse, inducing infectious anemia; im- provement after 2 to 4 years. In swine, experimentally, sometimes no outward and obvious signs of disease but blood, abnormal and infective ; sometimes severe; anemia, fever, prostration, loss of appe- tite. Thermal inactivation : At 58 to 60° C in 1 hour. Filterability : Passes Berkefeld Y filter candle. Other properties : Yiable in. blood in citrate saline at — 2° C for at least a year. Drying does not inactivate in 10 days but does in 1 month. Literature : DeKock., Union of South Africa, Dept. of Agv., 9th and 10th Re- ports for 1923, Pretoria 1924, 253-313; Habersang, Mouatshefte fiir prakt. Tier- heilk., SO, 1920, X71-176; Kutsche, ibid., 30, 1920, 557-56^; Peters, Jour. Am. Vet. FAMILY TRIFURACEAE 1283 Med. Assoc, 66, 1924, 363-366; Theiler and Kehoe, Union of South Africa, Dept. of Agr., 3rd and 4th Reports of the Direc- tor of Veterinary Research, 1915, 215-289. 2. Trifur gallinarum spec. nov. From Latin gallina, hen. Common name : Fowl -leucosis virus. Host : Gallus gallus (L.), chicken. Geographical distribution : United States, England, Europe. Induced disease : In chicken, neuro- lymphomatosis, with eye lesions (slate gray or bluish color replacing normal bay color of iris), anemia, hemocytoblastosis, lymphoid, erythroid or myeloid types of leucosis; the hemocytoblastosis is fol- lowed by infiltration of the central ner- vous system, peripheral nerves, iris, and many visceral organs by hemocytoblasts and lymphocytes, producing lesions some- times resembling neoplasms and consist- ing chiefly of hemocytoblasts (hemocyto- blastomata) ; marrow of radius and ulna becomes hyperplastic ; virus in blood plasma, blood cells, emulsions of organs; blood normal in its hydrogen-ion concen- tration ; recovery never complete ; ^ome stocks less susceptible than others. Transmission : By pen contact or con- taminated litter. Experimentally by in- travenous injection of cell -free filtrates. Not by the mosquitoes, Culex pipiens and Aedes aegypti {CULICIDAE) . Day- old chicks from iritis parents contain the infective agent and show some form of the induced disease in 80 per cent of the progeny if both parents show iritis, in 70 per cent if male is normal, 15 per cent if female is normal. Serological relationships : Specific neu- tralizing antibodies are formed in the rabbit as a result of injecting infective materials partly purified by sedimenta- tion in the ultracentrifuge. Thermal inactivation : At 56° C in 30 minutes. Filterability : Passes Berkefeld V, N, and W filter candles ; 1.5 per cent, but not often 3 per cent, collodion membranes; Seitz asbestos filter. Other properties : Viable after drying at least 54 days, in glycerine at least 104 days, at 4° C at least 14 days, at —60° C at least 6 months ; after freezing and thaw- ing, and after freezing in liquid air. Not viable after 14 days at 37.5° C. Particle diameter between 100 and 400 millimi- crons. Literature : Ellermann and Bang, Cent, f. Bakt., I Abt., Orig., 46, 1908, 4-5, 595- 609; Furth, Proc. Soc. Exp. Biol, and Med., 27, 1929, 155-157 ; Jour. Exp. Med., 53, 1931, 243-267; 55, 1932, 465-478, 495- 504; 58, 1933, 253-275; 59, 1934, 501-517; Furth and Miller, ibid., 55, 1932, 479-493 ; Hall et al., Am. Jour. Vet. Res., ;?, 1941, 272-279; J^rmai, Arch, wissensch. u. prakt. Tierhielk., 62, 1930, 113-131; Johnson, Virginia Agr. Exp. Sta. Tech. Bull. 56, 1934, 1-32; Johnson and Bell, Jour. Inf. Dis., 58, 1936, 342-348; Kabat and Furth, Jour. Exp. Med., 71, 1940, 55-70; 74, 1941, 257-261 ; Lee and Wilcke, Am. Jour. Vet. Res., 2, 1941, 292-294; Lee et al., Jour. Infect. Dis., 61, 1937, 1-20; Pierce, Am. Jour. Path., 18, 1942, 1127-1139; Ratcliffe and Stubbs, Jour. Inf. Dis., 56, 1935, 301-304. 1284 MANUAL OF DETERMINATIVE BACTERIOLOGY FAMILY VI. RABULACEAE FAM. NOV. Viruses of the Mumps Group, characterized in general by a special affinity for ( issues of the salivary glands. There is a single genus, Genus I. Rabula gen. nov. With characters of the family. Generic name from Latin rabula, pettifogger. The type species is Rabula inflans spec. nov. Key to species of the genus Rabula. I. Affecting man. 1. Rabula inflans. II. Affecting guinea pig. 2. Rabula Levis. III. Affecting hamster. 3. Rabula innocuus. IV. Affecting rat. 4. Rabula exiguus. V. Affecting mouse. 5. Rabula latens. 1. Rabula inflans spec. nov. From Latin inflare, to puff up. Common names : Mumps virus, virus of epidemic parotitis. Hosts : HOM I N I D AE—Homo sapiens L., man. Experimentally, also CERCO- PITHECIDAE—Macaca mulatta (Zim- mermann), rhesus monkeJ^ FELIDAE — Felis caius L., domestic cat. Geographical distribution : World-wide. Induced disease : In man, in order of frequency, parotitis, orchitis, meningo- encephalitis, pancreatitis, or ovaritis; rarely fatal ; when parotitis occurs, onset is sudden, with pain in one or both parotid glands, sometimes also with involvement of submaxillary and sublingual glands, swelling and malaise gradually disappear- ing within a week or 10 days ; there is virus in saliva 48 hours after onset ; orchi- tis, less common, is usually unilateral and may be accompanied by some epididymi- tis. In rhesus monkej', experimentally, acute, non-suppurative parotitis; focal necrosis in acinar epithelial cells of parotid gland, and secondary inflamma- tion; dissemination of lesions within the gland, enlargement of gland to palpation and pitting edema of jowl 6 to 8 days after inoculation, often with a rise of tempera- ture; cytoplasmic inclusion bodies in affected glands, staining pink, round or oval, 3 to 10 microns in diameter, often vacuolate, usually surrounded by a nar- row clear zone in the cytoplasm; blood and uninoculated salivary gland of af- fected animal not effective sources of virus. Transmission : Probably by droplets arising directly from infected individuals. E.xperimentally, by injecting sterile fluids containing virus into Stenson's duct of parotid gland in Macaca mulatta. Serological relationships : A specific complement -fixing antibody occurs in human and monkey convalescent serum and is demonstrable by the use of mon- key-gland antigen. Immunological relationships : Specific immunity induced by attack; passive immunization rarely successful. Thermal inactivation : At 55° C in 1 hour. Filterability : Passes Berkefeld V and N filter candles. Other properties : Viable in 50 per cent glycerine at 2° C at least 5 weeks, in 50 per cent glycerine at 10° C. at least 7 weeks, dried while frozen at least 7 weeks, in frozen saliva at least 3 weeks. FAMILY RABULACEAE 1285 Literature: Bloch, Am. Jour. Path., 13, 1937, 939-944; Enders and Cohen, Proc. Soc. Exp. Biol, and Med., 50, 1942, 180-184; Findlay and Clarke, Brit. Jour. Exp. Patli., 15, 1934, 309-313; .Johnson and Goodpasture, Jour. Exp. Med., 59, 1934, 1-19; Am. Jour. Hyg., 21, 1935, 46-57; ^3, 1936, 329-339 ; Am. Jour. Path., 12, 1936, 495-510. 2. Rabula levis spec. iiov. From Latin levis., trifling. Common name : Guinea-pig salivary- gland virus. Host: CAVIIDAE—Cavia porcellvs (L.), guinea pig (only known host; fetus more susceptible than post-natal animal, even if from immune mother) . Insusceptible species : Rabbit, rat, cat, chicken, pigeon, dog, mouse, monkey (Macacus rhesus). Geographical distribution: L'nited States, England. Induced disease : In guinea pig, sub- maxillary glands show swollen epithelial cells containing relatively dense acido- philic inclusions of granular material within enlarged nuclei, especially in ducts of the serous portion of the gland, and larger but fewer intracytoplasmic inclu- sions; experimentally, by intracerebral injection of young guinea pig, prodromal period of about 2 days, then elevation of temperature to 105 or 106° F ; a day later, hair raised, animal quiet; subsequently, irritability with tremors and slight con- vulsive movements ; by fifth day, usually prostration, jerking movements, and en- suing death ; brain shows no gross lesions but exudate over surface ; in meningeal exudate, many cells each containing an acidophilic mass within its nucleus; by subcutaneous injection, virus recoverable after 2 weeks from submaxillary glands, cervical lymph nodes, kidney, and lung, not from blood, liver, or spleen. Transmission : Experimentally, by in- oculation of submaxillary gland or by intracerebral or subcutaneous injection of materials from infected glands; with difficulty from brain to brain. Pilocar- pine stimulation increases numbers of inclusions. Serological relationships : Specific neu- tralizing antibody is found in blood serum of animals that are carrying virus in their submaxillary glands. Immunological relationships: Active immunity may be dependent on existence of more or less active lesions. Thermal inactivation : At 54° C in 1 hour. Filterability : Passes Berkefeld N filter candle. Other properties : Viable in 50 per cent glycerine at least 11 days. Strains: An unusually virulent strain, killing infected animals whatever the route of injection, has been described but not given a distinctive name (Rosenbusch and Lucas, Am. .Tour. Path., 15, 19.39, 303-340). Literature : Andrewes, Brit. Jour. Exp. Path., 11, 1930, 23-34; Cole and Kuttner, Jour. Exp. Med., U, 1926, 855-873; Hud son and Markham, ibid., 55, 1932, 405-415 ; Jackson, Jour. Inf. Dis., 26, 1920, 347- 350; Kuttner, Jour. Exp. Med., 46, 1927, 935-956; Kuttner and T'ung, ibid., 62, 1935, 805-822; Lucas, Am. Jour. Path., 12, 1936, 933-948; Markham, ibid., U, 1938, 311-322; Markham and Hudson, ibid., 12, 1936, 175-182; Pearson, ibid., 6, 1930, 261-274; Scott, Jour. Exp. Med., 49, 1929, 229-236; Scott and Pruett, Am. Jour. Path., 6, 1930, 53-70. 3. Rabula innocuus spec. nov. From Latin innocuus, harmless. Common name : Hamster salivary- gland virus. Host : CRICETIDAE—Cricetulus gri- seus M. Edw., Chinese hamster. Insusceptible species: MURIDAE — rat; Mus musculus L., white mouse. Geographical distribution : China. Induced disease : In hamster, no obvi- ous disease externally but inclusion bodies in submaxillary glands. 1286 MANUAL OF DETERMINATIVE BACTERIOLOGY Thermal inactivation : At 56° C in 30 minutes. Literature : Kuttner and Wang, Jour. Exp. Med., 60, 1934, 773-791. 4. Rabula exiguus spec. nor. From Latin exiguus, petty. Common name : Rat salivary -gland virus. Host : M URIDAE— rat. Insusceptible species: M URIDAE — Mus musculus L., mouse. CRICETI- DAE — Cricetulus griseus M. Edvv., Chi- nese hamster. Geographical distribution : China, Canada. Induced disease : In rat, no obvious disease externally, but intranuclear in- clusions in cells of the submaxillary glands. Literature : Kuttner and Wang, Jour. Exp. Med., 60, 1934, 773-791 ; Thompson, Jour. Inf. Dis., 50, 1932, 162-170. 5. Rabula latens spec. nov. From Latin latens, hidden or lurking. Common name : Mouse salivary -gland virus. Host: M URIDAE — Mus muscvlus L., mouse. Insusceptible species: MURIDAE — rat. CRICET I DAE— Cricetulus griseus M. Edw., Chinese hamster. LEPOR- I DAE— rahhii. CA VIIDAE—Cavia porcellus (L.), guinea pig. Geographical distribution : China, Canada, United States. Induced disease : In mouse, no obvious disease externally, but inclusion bodies in acinar tissue of serous and mucous por- tions of submaxillary glands ; occasionally also in duct cells or alveolar cells of paro- tid gland; affected cells hypertrophied. In Swiss white mice, extensive lesions in liver and spleen but emulsions of these organs fail to infect; rare pancreatic lesions. Transmission : Experimentally, by in- traglandular, subcutaneous, intraperi- toneal, intra testicular or intracerebral inoculation; inclusion bodies appear in salivary glands irrespective of site of inoculation. Thermal inactivation : At 60° C in 30 minutes. Filterability : Passes Berkefeld V filter candle. Literature : Kuttner and Wang, Jour. Exp. Med., 60, 1934, 773-791 ; McCordock and Smith, ibid., 63, 1936, 30.3-310; Thompson, Jour. Inf. Dis., 58, 1936, 59-63 SUPPLEMENT NO. 3 PLEUROPNEUMONIA AND PLEUROPNEUMONIA-LIKE ORGANISMS (BORRELOMYCETACEAE) Louis Dienes Boston, Mass. May, 1945 THE ORGANISM OF CONTAGIOUS BOVINE PLEURO- PNEUMONIA AND RELATED ORGANISMS* INTRODUCTION The organism of bovine pleuropneumonia is similar in certain respects to filterable viruses. Both in infected tissue and in cultures, small elements are present which pass through filters that retain bacteria. The organism is not stained well by the usual bacterial stains and can be made visible only by using special methods. Bovine pleuropneumonia and other diseases caused by similar organisms were originally attributed to filterable viruses. These organisms are different from \'iruses in an important point; namely, they grow on suital^le media in the absence of living host cells. The cul- tures consist of pleomorphic elements, the nature of which has only slowly become apparent. By the studies of Nowak (Ann. Inst. Past., 43, 1929, 1330), Turner (Jour. Path, and Bact., 4i, 1935, 1) and Klieneberger and Smiles (Jour. Hyg., 4^, 1942, 110), it has been established that the pleo- morphic forms are part of a reproductive cycle different from binary fission. The small elements in the cultures swell up into large round forms which reproduce the small elements within their membranes. The mor- phology of the organism is further complicated by the fact that long branch- ing filaments are present in freshly isolated bovine strains. These break up into granules or parts of the filaments swell up into large round forms. In the judgment of some investigators, these properties, in addition to un- usual softness and fragility, exclude the organism of bovine pleuropneu- monia and similar organisms from the order of true bacteria. Ledingham (Jour. Path, and Bact., 37, 1933, 393) has classified them with the Actino- myces. Later, Turner (Jour. Path, and Bact., 4i, 1935, 1) placed them in an independent order, Borrelomycetales, while Sabin (Bact. Rev., 5, 1941, 58) has even placed them in an independent class, Paramycetes. The observations of the present author give support to the classification of Buchanan (Jour. Bact., 3, 1918, 44) who placed the genus Asterococcus Borrel et al. with the organism of bovine pleuropneumonia {Asterococcus mycoides Borrel et al.) as type, together with the genus Haemophilus Win- slow et al. in subtribe Haemophilinae Buchanan of the tribe Bacterieae Trevisan emend. Buchanan. In many strains of the pleuropneumonia *Common names have been used through Supplement No. 3 (except for Astero- coccus and A. rmjcoides) as the author believes that a more suitable nomenclature than any thus far proposed should be developed when agreement is reached as to the nature of these organisms. Specific names that have been proposed are given merely as a matter of record. No new names have been introduced. 1289 1290 MANUAL OF DETERMINATIVE BACTERIOLOGY group, the small forms appear in appropriate preparations as small bipolar stained bacilli. The transformation of the bacilli to round bodies of variable size often occurs in bacterial cultures and is not specific for the pleuropneumonia group. Furthermore, it has been observed in several species of bacteria that they reproduce in the large round forms in a manner similar to that observed in the pleuropneumonia group. Thus the form variation and reproductive processes observed in the pleuropneumonia group are not specific to this gi'oup. They represent general bacterial properties and should be included in the definition of the true bacteria. According to these considerations, the organisms belonging to the pleuropneumonia group are small, Gram-negative bacilli often showing bipolar staining and their distinctive characteristic is the tendency to swell up into round forms and multiply by the reproduction of l^acilli in the round forms. Their habitat is in the mucous membranes of animals and man and many of them are pathogenic. They are exacting in their media requirements and usually require fresh animal serum for their growth. These properties indicate a close similarity to the species now included in the genera Pasteurella and Haemophilus. The pleuropneumonia group might well be classified in the same or a closely related family. It is un- certain whether the strains isolated from earth and sewage should be classi- fied with the strains isolated from animals and men. The soil and sewage strains are less soft, stain more easily and gi-ow abundantlj'^ without animal serum. The strains isolated from bacterial cultures are most probably variant forms of the bacteria and should be classified with the parent organisms. The viruses of psittacosis and lymphogranuloma present similarities to the pleuropneumonia group both in morphology and in their methods of reproduction. This gives added weight to the thought that the pleuro- pneumonia group represents an intermediary stage in the evolution of the small. Gram-negative bacteria of the mucous membranes into the filterable viruses. BORRELO\n'CETACEAE 1291 I. THE PLEUROPNEUMONIA GROUP. (Borrclomijcetacene Turner, Jour. Path, and Bact., 4^ , 1035, 25; Pnrasitnceac Sabin, Bact. Rev., 5, 1041, 58.) The organisms are soft and fragile. Without special precautions the}' are often distorted or entireh' destroyed in microscopical preparations. The cultures contain pleomorphic elements: Small granules, bacilli, bacillary filaments and round forms varying in size from a few tenths of a micron to 10 microns or more. Autolyzed round forms may coalesce into large empty blebs. The round forms are part of a reproductive cycle. They are produced by the swelling of the bacillary forms and filaments and reproduce granules or filaments by inside segmentation or multiple germination. In freshly isolated bovine strains, the filaments show apparent or true branching and reproduce the small forms by segmentation. The smallest growing units may not be larger than .15 to .28 micron and pass through filters that retain bacteria. On agar, tiny colonies (0.1 to 0.6 mm) develop in great numbers. The colonies invade the agar and after 2 to 5 days growth have an opaque center embedded in the agar and a thin peripheral zone. The surface has a rugged or granular appearance due to the develop- ment and autolysis of the large forms. After a few days growth, the cultures usualy show pronounced autolysis. The parasitic strains require fresh animal serum for growth. There is a single genus. Genus I. Asterococcus Barrel et al. (Borrel, Dujardin-Beaumetz, Jeantet and Jouan, Ann. Inst. Past., 2^, 1910, 179; Coccobacillus Martzinovski, Ann. Inst. Past., 35, 1911, 91 ; Microtnyces Frosch, Arch, f. wissensch. u. prakt. Tierheilk., 49, 1923, 35 and 273; not Micromyces Dangeard, Le Botaniste, 1, 1888, 55; Mycoplasma Nowak, Ann. Inst. Past., 43, 1929, 1349; Aster- omyces Wroblewski, Ann. Inst. Past., 47, 1931, 105; Borrelomyces Turner, Jour. Path, and Bact., 4^ > 1935, 25; Bovimyces Sabin, Bact. Rev., 5, 1941, 57.) Characters as for the family. The type species is Asterococcus mycoides Borrel et al. 1 . Asterococcus mycoides Borrel et al . agar cultures correspond with the descrip- (Le microbe de la peripneumonia, Nocard tion given for the group, and Roux, Ann. Inst. Past., 12, 1898, 240; Broth cultures are slightly opalescent Borrel, Dujardin-Beaumetz, Jeantet and and, upon shaking, the cultures of fresh Jouan, Ann. Inst. Past., 24, 1910, 168; strains exhibit silk-like whorls, due to Coccobacillus mycoides periyneumoniae the presence of long chains and filaments. Martzinovski, Ann. Inst. Past., 25, 1911, The cultures after prolonged incubation 914; Micromyces peripneumoniae bonis consist of small granules, con/agffosae Frosch, Arch. f. wissensch. u. Biochemical activity: Old colonies on prakt. Tierheilk., 49, 1923, 35 and 273; serum agar develop a brownish color. Mycoplasma peripneumoniae Nowak, Freshly isolated strains reduce hemo- Ann. Inst. Past., 4S, 1929, 1530; Astero- globin. Glucose, fructose, mannose, mal- myces peripneumoniae bovis Wroblewski, tose, and dextrin are fermented with the Ann. Inst. Past., ^7, 1931, 94; Borrelo- production of acid but no gas. The cul- niyces peripneumoniae Turner, Jour. tures are bile soluble. Path. Bact., 4-', 1935, 1 ; Boyimyces piei/- The strains isolated from cattle are ropneumoniae Sabin, Bact. Rev., 5, 1941, homogeneous in serological reactions and 57.) distinct from the other members of the Morphologj' of cells and appearance of group. 1292 MANUAL OF DETERMINATIVE BACTERIOLOGY Habitat : It is the causative agent of contagious bovine pleuropneumonia. The disease can be transferred to sheep, goats and water buffaloes, but not to mice, rats, rabbits or other experimental animals. 2. The organism of agalactia of sheep and goats. (Le microbe d'agalaxie con- tagieuse, Bridr4 and Donatien, Ann. Inst. Past., 39, 1925, 925; Anulomyces agalaxiae Wroblewski, Ann. Inst. Past., 47, 1931, 111; Borrelomyces agalactiae Turner, Jour. Path, and Bact., 41, 1935, 25; Capromyces agalactiae Sabin, Bact. Rev., 5, 1941, 57.) These organisms are very similar to the former organisms in morphology, appear- ance of the cultures and growth require- ments. Usually the growth is less vigor- ous, the colonies remain smaller, and the elements of the cultures are more delicate and less easily visible than those of bovine pleuropneumonia. Characteris- tic crystals develop in the cultures. Serologically and immunologically this species is distinct from the bovine spe- cies. It is the cause of a systemic disease in sheep and goats with involvement of the joints, eyes, and, in lactating animals, the mammary glands. Other species are not susceptible. 3. Pleuropneumonia-like organisms in dogs. {Asterococcus canis, Types I and II, Schoentensack, Kitasato Arch. Exp. Med., 11, 1904, 227; 13, 1936, 175; Cano- myces ■pulmonis I and II, Sabin, Bact. Rev., 5, 1941, 57.) Both types produce slight uniform opalescence in broth. Type I grows in granules and coarse colonies and is ap- parently pathogenic for dogs. Type II grows in somewhat larger granular colonies. They are serologically distinct from each other and from the other members of the pleuropneumonia group. The connection of these organisms with distemper is not proven. 4. Pleuropneumonia-like organisms in rats. Ls (Klieneberger and Steabben, Jour. Hyg., 37, 1937, 143; Jour. Hyg., 40, 1940, 223; Murimyces pulmonis Sabin, Bact. Rev., 5, 1941, 57.) L4 (Klieneberger, Jour. Hyg., 38,1938, 458; Murimyces arthritidis Sabin, loc. cit.) The pyogenic virus of Woglom and Warren (Jour. Exp. Med., 68, 1938, 513) and L7 of Findlay, MacKenzie, Mac- Callum and Kleineberger (Lancet, 237, 1939, 7) are identical with L4. The organisms isolated from infected joints by Beeuwkes and Collier (Jour. Inf. Dis., 70, 1942, 1) and Preston {ibid., 70, 1942, 180) probably are identical with L4 but they were not typed serologically. The requirements for growth, the ap- pearance of colonies and the morphology are very similar to those of the type strain with the difference that long fila- ments are not observed either in liquid or solid media. The strains isolated from rats belong to two serological types. L3 was cultivated from chronic lung abscesses, but the num- ber of strains typed is not sufficient to ascertain that all strains isolated from this source belong to one type. The L3 strains are not pathogenic for rats in artificial infection. They produce sup- puration in mice when they are injected with agar. L4 which is serologically different from Ls was cultivated from abscesses and spontaneous polyarthritis. It produces polyarthritis both in mice and rats. It is not infectious in monkeys, rabbits and guinea pigs. Both L3 and L4 were recov- ered from the brains of mice kept in the same room with rats. According to Klieneberger, L3 usually produces somewhat larger and coarser colonies than L4; L3 grows in broth in small granules, while L4 produces an opalescent growth. BORRELOMYCETACEAE 1293 5. Pleuropneumonis-like organisms in mice. (Sabin, Science, 88, 1938, 575, and Bact. Rev., 5, 1941, 1; Findlay, Kliene- berger, MacCallum and MacKenzie, Lan- cet,^35, 1938, 1511.) A strain isolated by Sullivan and Dienes (Proc. Soc. Exp. Biol, and Med., 41, 1939, 620) is identical with Type A. Five groups of strains, distinct sero- logically, and, to a certain extent, distinct also in their pathological properties, have been isolated from mice. These are types A, B, C, D, and E of Sabin. The strains are closely similar to each other and to the rat strains. It is questionable whether the slight differences in the ap- pearance of the colonies and in the mor- phology of the cultures are of significance. Type A (Musculomyces neurolyticus Sabin) is usually present in the conjunc- tiva and was i.solated also from the lung and brain. Intercerebral injection of Type A produces in mice a characteristic rolling disease due to a toxin which is also present in broth cultures. Intravenous injection produces a transient polyarthri- tis without damage to the cartilage or ankylosis. Type A is serologically simi- lar to Lo of Klienebergcr, (.lour. liyg., 40, 1940, 204). Type B {Musculomyces arthrotropicus Sabin) was isolated from the brain and from the nasal mucosa. It produces no rolling disease and no soluble toxin. In mice, intravenous injection usually pro- duces a chronic arthritis often leading to ankylosis. Types C, D and E {Musculomyces his- totropicus Sabin) were isolated from the same location as Type B and produce similar arthritic lesions. They are sero- logically distinct from Type B and from each other (Sabin, Science, 90, 1939, 18 and Sabin and Johnson, Proc. Soc. Exp. Biol, and Med., 44, 1940, 569). Le isolated from mice by Findlay et al. (Trans. Roy. Soc. Trop. Med. Hyg., 33, 1939-40, 6) and the strains of Edward (Jour. Path, and Bact., ,50, 1940, 409) were not compared serologically with the types of Sabin. 6. Pleuropneumonia-like organisms in man. (Dienes, Proc. Soc. Exp. Biol, and Med., 44, 1940,468; Beveridge, Med. Jour, of Australia, 30, 1943, 479; Kleine- berger, Lancet, 2, 1945, 46.) They are present in about 30 per cent of women in the genitals and they were isolated from suppurative processes orig- inating from this source. In men they were found in urethritis, cystitis and chronic prostatitis. The appearance of the colony, the mor- phology and growth i-equirements cor- respond with the animal strains. The human strains grow less abundantly in serum broth than the animal strains. One strain was found by Sabin (Proc. Soc. Exp. Biol, and .Med., 44, 1940, 569) to be serologically different from the strains isolated from rats and mice. It is not known whether the strains are sero- logically uniform. There is a slight vari- ation in colon}^ form, in the tendency to grow in filaments, and in the abundance of growth, but the variation between the strains is less than the variation due to .slightly different cultural conditions. Mice and rats are usually not suscepti- ble to infection with the human strains; however, several A^oung mice from a single litter were killed in three to six days by subcutaneous or intraperitoneal injection of one strain. 7. Pleuropneumonia-like organisms in chick embryos. (Van Herick and Eaton, Jour. Bact., 50, 1945, 47.) Organisms have been isolated from chick enibryos which conform to the pleuropneumonia group with regard to morphology, the appearance of colonies on agar and filterabilit}'. The cultures agglutinated red blood cells from various animals. The relation of this strain to the coccobacillary bodies of Nelson (see Section II) has not been studied. 1294 MANUAL OF DETERMINATIVE BACTERIOLOGY II. ORGANISMS OF UNCERTAIN CLASSIFICATION. Similar to the Pleuropneumonia Group. 1. Coccobacillary bodies of Nelson. (Nelson, Science, 82, 1935, 43; Jour. Exp. Med., 65, 1937, 833; Jour. Exp. Med., 72, 1940, 645.) Nelson isolated a small bacillary organ- ism apparently connected with coryza and infectious catarrh from the nasal passages of fowls and from the nasal passages and the middle ear of mice and rats. Their size appeared to be 0.3 to 0.4 micron in microscopical preparations and they passed through a filter with a pore size of 640 millimicrons. They were isolated in tissue cultures but they grow also in the cell -free and heated supernatant fluid. The freshly isolated cultures did not grow on blood or on artificial media; however, after 120 passages in tissue cul- tures the fowl coryza bodies grew on blood agar slants. On ascitic agar this strain forms colonies very similar to those of the pleuropneumonia group with a dark center surrounded by a thin periphery. The organisms in the top layer are sometimes considerablj^ enlarged, but no web-like structure is produced. The organism is less soft and the individual organisms maintained their form in the preparations as do bacteria, and the ten- dency to grow into the agar is less pro- nounced than in the pleuropneumonia group. The organism isolated from rats is more pleomorphic than the others. The coccobacillary bodies were not studied with methods appropriate to de- termine whether they belong to the pleu- ropneumonia group and whether the mouse and rat strains are identical with the pleuropneumonia-like organisms iso- lated from mice and rats. 2. Filterable organisms from sewage and soil. (Fam. Saprophytaceae Sabin, Genus Sapromi/ces Sabin, Bact. Rev., 5, 1941, 59.) The strains isolated by Laidlaw and Elford (Proc. Roy. Soc. London, B, 120, 1936, 292 ; Sapromyces laidlawi A B and C , Sabin, loc. cit.) and Seiffert (Cent. f. Bakt., I Abt., Orig., 139, 1937, 337) ac- cording to 0rskov (Cent, f . Bakt., I Abt., Orig., HI, 1938, 230) and Klieneberger (Jour. Hyg., 40, 1940, 204) are closely similar to the organisms of the pleuro- pneumonia group. They are filterable, and the smallest reproductive units of those which we have appropriately ex- amined were found to be between .125 and .175 micron. The colonies are simi- lar in appearance to the colonies of the pleuropneumonia group. The broth cultures consist of granules and round globular elements ; the surface layer of agar colonies sometimes swells up into large round forms. They grow without serum, but small amounts of serum accelerate the growth. They grow both at 30° and at 37°C. and remain alive in cultures kept cold for several months. The broth cultures grow abun- dantly with a strong opalescence or sedi- ment . vSerologically the strains are dis- tinct from the other members of the pleuropneumonia group and all but one are more or less similar to each other. III. PLEUROPNEUMONIA-LIKE ORGANISMS ISOLATED FROM BACTERIAL CULTURES. 1. Pleuropneumonia-like organisms isolated from Streptobacillus monili- formis. Li (Klieneberger, Jour. Path, and Bact., 40, 1930, 93; Jour. Hyg., 42, 1942, 485; Dienes, Jour. Inf. Dis., 65, 1939, 24; Jour. Bact., U, 1942, 37; Mus- culomyces streptobacilli-moniliformis Sa- bin, Bact. Rev., 5, 1941, 57; Heilman, Jour. Inf. Dis., 69, 1941, 32; Brown and Nunemaker, Bull. Johns Hopkins Hosp., 70,1942,201.) Cultures isolated from different strains BORRELOlVTiCETACEAE 1295 of Streptobacillus moniliformis vary con- siderably in the appearance of the colonies, the tendency to reversion to bacillarj' form, and the degree of autoly- sis. The colonies are considerably larger than the colonies of the human or animal pleuropneumonia-like strains; they may reach 1 to 2 mm. Usually a wide periph- eral zone is present and development and autolysis of the large bodies produces a coarse appearance in the colonies. Sometimes no peripheral zone develops, the colony is dome -shaped, and the large bodies have no tendency to autolyze. The young colonies (twelve hours incuba- tion) grow into the agar as loose strands of more or less swollen granules. Serum broth cultures grow in small clumps usu- ally adhering to the wall of the test tube. The cultures consist of small granules, small polar-staining bacilli and diph- theroid-like forms which swell to large round forms. In the top layer of fully developed colonies, the well-stained large bodies may be as large as 10 to 20 microns. By vacuolization they transform into empty blebs. By segmentation of their contents, the large forms may reproduce the small bacillary forms. In suitable preparations chromatin bodies are visible both in the small and large forms. The small forms are filterable through Berk- feld candles; the size of the smallest particles has not been exactly deter- mined. The organism is very soft and fragile . Their growth requirements and bio- chemical activities are similar to those of Streplohacill us moyiiliform is . Growth occurs on nutrient agar con- taining animal .serum or egg yolk. Some- times there is a slight growth on boiled blood agar plates without serum. Good growth is obtained in a mineral solution with 0.1 per cent starch. Growth is both aerobic and anaerobic. The Li form is more resistant to heat and to aging of the culture than is the streptobacillus and it has a remarkable resistance to penicillin to which the bac- teria are very sensitive. Like the bacil- lus, Li produces acid but no gas from glucose, maltose, fructose, salicin, starch, and dextrin. It gives no oxidase test. Serologically the Li form is similar to Streptobacillus moniliformis and different from the members of the pleuropneu- monia group. It has no pathological effect on mice, rats or guinea pigs. It does not produce an infection of the chicken embryo. It can be isolated from freshly isolated strains of Streptobacillus moniliformis, from several-day old broth and agar cultures, from broth cultures heated at 56°C. and usually also from 48 hour agar cultures if they are incubated at 28° to 30°C. It is questionable whether the Li form has been isolated directly from rats. Klieneberger (Jour. Uyg., 40, 1940, 204) isolated a similar strain from a bacterium similar to Streptobacillus moniliformis which caused abscesses in guinea pigs. Whether this bacterium was identical or different from Streptobacillus monili- formis was not determined. 2, Pleuropneumonia-like organisms isolated from Bacteroides funduliformis. Dienes (Proc. Soc. Exp. Biol, and Med., 47, 1941, 385) and Dienes and Smith (Jour. Bact., 48, 1944, 125) isolated cul- tures from two strains of Bacteroides funduliformis which could be propagated indefinitely and which in morphology and in the appearance of colonies were closely similar to Li. The young colonies consisted of similar strands of granules growing into the medium. The surface of fully developed colonies consisted of large bodies and a honey-comb-like structure. The well isolated colonies grew usually to a fairly large size (1 to 2 mm). Both strains, transplanted every two or three days through several months, failed to reproduce bacteria either on agar or submerged in broth. No growth was obtained in liquid cul- tures. 1296 MANUAL OF DETERMINATIVE BACTERIOLOGY The strains, like the parent organism, are strictly anaerobic and the cultures have the characteristic odor of the parent strain. It was observed in slide cultures that the L type of colonies develop from large round forms which were produced in the cultures of the parent organism by grad- ual swelling of the bacteria. In cultures of eight pleomorphic strains of Bacteroides, the L type of colonies developed in three strains under ap- propriate conditions. The bacteria swelled into large round bodies in all eight strains. The serological properties of the L strains have not thus far been studied. Neither the parent organisms nor the L type strains had any pathological effect on laboratory animals. 3. Pleuropneumonia-like organisms in a species of Flavobacterium. (Dienes, Jour. Bact., U, 19-i2, 37.) Tiny colonies entirely similar in ap- pearance to young Li colonies were iso- lated from the cultures of a species of Flavobacterium . The bacterium when freshly isolated produced two types of colonies on blood agar plate ; large colonies consisting of small regular bacilli and tiny colonies in which the bacteria became pleomorphic and swelled up to form large round bodies. The tiny colonies after 48 hours of incuba- tion became autolyzed, and one or several L type of colonies started to grow under them. These colonies could be trans- planted and gave abundant growth for two generations, but always died out in the third. Bacterial forms were not reproduced either on agar or in broth. The L type of growth was not patho- genic for mice though the parent organism was highly virulent. 4. Development of tiny colonies in other bacteria. The development of tiny colonies simi- lar in appearance to young colonies of the pleuropneumonia group has been ob- served in cultures of Escherichia coli, Haemophilus influenzae, and Neisseria gonorrhoea (Dienes, Jour. Bact., 44, 1942, 37; Proc. Soc. Exp. Biol, and Med., 44, 1940, 476). In all cases preceding their development, the organisms ot the parent strains swelled into large round bodies, and in Escherichia coli and Haemophilus influenzae the development of the L type of colonies from these large forms was observed. Thus far these tiny colonies have not been isolated in pure cultures. INDEX* SOURCES AND HABITATS (All references to viruses will be found under the heading fViruses) Abnormal Milk, see Dairy Products Agar Bacillus, 734 Containing iodiform Micrococcus, 272 Digesting bacteria Acetobacier, 692 Achromobacter, 628 Agarhacterium, 628, 629, 630 Bacterium, 625, 626 Flavobacferium , 631 Pseudomonas, 177, 178, 697, 698, 700 Vibrio, 200, 203, 204, 702, 703 Air, also see Dust Actiyiomijces, 968, 969, 970, 972, 973 Bacillus, 647, 648, 649, 650, 651, 653, 654, 655, 658, 659, 661, 663, 664, 665, 667, 668, 669, 670, 671, 672, 738, 741, 742, 743, 744, 747, 748, 749, 750, 751, 752, 754, 756, 758 Bacterium, 602, 643, 672, 674, 676, 678, 680, 684, 761 Chromobacterivm, 232, 233 Clostridium, 803 Corynebacierium , 386 Flavobacterium , 611 Gaffkya, 283 Leuconostoc, 348 Micrococcus, 237, 239, 244, 251, 252, 253, 255, 256, 260, 261, 268, 271, 272, 273, 276, 278. 281 Nocardia, 915, 975 Pacinia, 696 Planococcus, 281 Pseudomonas, 95, 96, 147, 149, 174 Rhodococcus, 281 Sarcina, 288, 290 Staphylococcus , 282 Streptococcus, 336, 338, 340 Streptomyces, 935, 969 Strepfothrix, 975 Air — brewery Sarcina — contamination Nocardia, 915 Planococcus , 281 Pseudomonas, 174 Staphylococcus, 282 Contamination on cooked jxjtato Chr om abaci eriu7n, 232 Amphibia, Common Names African toads Vibrio, 197 Frogs Bartonella, 1108 Eberthella, 534 Flavobacterium , 439 Malleomyces, 556 Micrococcus, 281 Mycobacterium, 883, 884, 885, 890 Pseudomonas, 102, 103 Salmonella, 532 — , abscesses Micrococcus, 281 — , feces Bacillus, 742 — , intestine Chitin-digesting bacteria, 632 Spirochaeta, 1065, 1069, 1070 — , large intestine Bacillus, 742, 754 Spirillum, 217 — , wound infections Eberthella, 534 Salamander Pseudomonas, 102 * Prepared by Prof. Robert S. Breed and Mrs. Margaret E. Breed, Geneva, New York, August, 1947. t Prepared by Frances O. Holmes, Rockefeller Institute for Medical Research, Princeton, New Jersey, July, 1947. 1297 1298 INDEX OF SOURCES AND HABITATS Amphibia {continued) Toads Arlhrornitis, 1003 Bacillus, 744, 754 Micrococcus, 281 Serratia, 462 Vibrio, 197 ■ — , abscesses Micrococcus, 281 — , intestine Spirochaeta, 1066 — , large intestine Spirillum, 217 Treponema, 1075 — , rectum Bacterium , 760 — , tadpoles Bacillus, 742 Tree toads Spirochaeta, 1066 Amphibia, Diseases of Frogs Pseudotuberculosis Malleomyces, 556 Tuberculosis Mycobacterium , 883, 884, 885, 8'JO Red leg Pseudomonas, 102, 103 Amphibia, Scientific Names Alijtes sp., 742 Bufo americanus, 462, 742, 754, 760. 1066, 1075 Hyla sepientrionulis, 1066 Leptodactylus ocellalus, 1108 Leptodactylus pentadactylus, 1113 Rana pipiens, 462 Rana tamporaria, 742, 754, 1065, 1069, 1070 Xenopus luevis, 439 Animal Diseases (Vertebrate), also see Amphibia, Birds, Fishes, Reptiles, and Mammals Vertebrates Cold-blooded Mycobacterium , 887 Warm-blooded ' Neisseria, 300 Rasteurella, 548, 549 Animal Products (Vertebrate; Catgut for sutures Bacillus, 815 Gelatin Colony on old plate Micrococcus, 281 Containing iodoform Micrococcus, 272 Spoiled Bacillus, 756 Glue Black discoloration Chromobacterium , 233 Hides, salted Pseudomonas, 110 Sarcina, 289 Animal Sources and Diseases (Inverte- brate) also see Arthropoda, Insecta, Mollusca, and Protozoa Animal Sources (Invertebrate), Com- mon Names Annelid, Marine Crisiispira, 1057 Blood suckers Bacillus, 746 Bacterium, 679 Echinoderm Crist ispira, 1056 Leeches Bacillus, 746 Bacterium , 679 Tunicate Spirochaeta, 1066 Animal Sources (Invertebrate), Scien- tific Names Asterias rubens, 1056 Caesira retortiformis , 1066 Hirudo spp., 679, 746 Polydora flava, 1057 Animal Sources (Vertebrate), also see Amphibia, Birds, Fishes, Reptiles, and Mammals Vertebrates, Intestine Aerobacter, 455 Escherichia, 447, 449, 450 Paracolobactrum , 460 A'ertebrates, Warm-blooded Salmonella, 503, 510, 517, 528 Vibrio, 196 INDEX OF SOURCES AND HABITATS 1299 Arthropoda, Common Names Arachnids Rickettsia, 1098, 1099 Crustacea Crayfish, intestine Chitin-digesting bacteria, 632 Fresh water Bacterium, 678 Paste uria, 836 Horseshoe crab, shell Bacterium, 632 Marine Bacterium, 635 Small Crustacea Eubacterium, 367 Paste uria, 836 Myriapoda Diplopods F us if or mis, 694 Millipeds, intestine Arthromitis, 1003 Alites Rickettsia, 1098 Haemohartonelln , llOo Bird mites Rickettsia, 1091 Human mites Rickettsia, 1091, 1092 Rodent mites Rickettsia, 1091, 1092 Spiders Rickettsia, 1098 Ticks Borrelia, 1059 Haemobartonella, 1105 Klebsiella, 459 Rickettsia, 1097, 1098, 1099 Spirochaeta, 1068 Bont ticks, Rickettsia, 1089, 1094 Dog ticks, 1088, 1096, 1098 Rickettsia, 1088, 1089, 1096, 1098 Rabbit ticks Rickettsia, 1088 Sheep ticks Rickettsia, 1097 Wood ticks Rickettsia, 1088, 1093, 1096 Arthropoda, Scientific Names Crustacea Daphnia sp., 836 Gammarus zschokkei, 678 Limulus polyphemus, 632 Talorchcstia sp., Ill Millipeds Julus marginatus, 1003 Mites Allodermanyssus sanguineus, 1092 Tromhicula akamushi, 1091 Trombicula deliencis, 1091 Trombicula flelcheri, 1091 Trombicula walchi, 1091 Ticks Amblyomma americanum, 1088, 1093, 1098 Amblyomma brasiliensis, 1088 Amblyomma cajennense, 1088 Amblyomma hebraeum, 1089, 1094 Amblyomma maculatum, 1098 Amblyomma spp., 1088 Amblyomma striatum, 1088 Amblyomma variegatum, 1094 Boophilus decoloratus, 1089 Dermacentor albipictus, 459 Dermacentor andersoni, 1088, 1093, 1096, 1098 Dermacentor occidentalis, 1093 Dermacentor spp., 1088 Dermacentor variabilis, 1088 Haemophysalis humerosa, 1093 Haemophysalis leachi, 1089 Haemophysalis leporis-palustris, 1088, 1093 Haemophysalis spp., 1088 Hyalomma spp., 1095 /xodes dentatus, 1088, 1093 Melophagus ovinus, 1068, 1097 Ornithodoros erraticus, 1066 Orniihodoros hermsi, 1064 Ornithodoros latiorensis, 1069 Ornithodoros marocanus, 1067 Ornithodoros moubata, 1060 Ornithodoros normandi, 1068 Ornithodoros parkeri, 1064 Ornithodoros rudis, 1064 Ornithodoros spp., 1063, 1088 Ornithodoros tholozani, 1060 Ornithodoros turicala, 1064 Ornithodoros venezuelensis, 1064 Rhipicephalus appendicidatus, 1089 Rhipicephalus bursa, 1097 Rhipicephalus decoloratus, 1062 1300 INDEX OF SOURCES AND HABITATS Arthropoda (continued) Ticks (continued) Rhipicephahis sanguineus, 1089, 1096, 1098 Rhipicephalus spp., 1088 Arthropod Vectors, see Arthropoda, and Insecta Bacterial Cultures Pleuropneumonia-like organisms, iso- lated from Bacleroides fundidifonnis, 1295 Flavohacteiium sp., 1296 Streptobacilius moniliformis, 1294 Beer Acetohacter, 183, 184, 185, 186, 188, 189 Bacterium, 680 Flavohacterium , 441 Pediococcus, 249, 250 Pseudomonas, 94, 176 Bottled Micrococcus, 260 Double Acetohacter, 183 Ginger Bacterium, 362 Ropy Acetohacter, 188, 189 Streptococcus, 250 Sarcina-sick Pediococcus, 249 Spoiled Achromobacter, 423 Lactobacillus , 360 Pediococcus, 249, 250 Streptococcus, 345 Beermash Spoiled Pediococcus, 249, 250, 260 Beer Wort Acetohacter, 183, 185, 186 Bacillus, 758 Inflabilis, 823 Lactohacterium, 363, 364 Pseudomonas, 146 Birds, common names Bullfinch Hicketlsia, 1095 Canaries Bacillus, 530 Pasteurella, 554 Shigella, 540 Chaffinch, stomach and intestine Micrococcus, 260, 266 Coot, stomach Micrococcus, 260, 263 Crow, stomach contents Micrococcus, 269 Cuckoo, throat Corynelacterium , 403 Dove, intestine Bacillus, 666 Bacterium, 760, 761, 762 Micrococcus, 252, 254, 270, 271, 274, 276 Sarcina, 292 Dove, stomach Bacterium, 760 Micrococcus, 252, 254, 270, 271, 274, 276 Duck Borrelia, 1059 Pasteurella, 552, 554 Pfeifferella, 554 Salmonella, 505, 518, 523, 527 Duck, skin Corynebacterium, 406 Finch, intestine Micrococcus, 257, 259 — , stomach Bacterium, 761 Micrococcus, 251, 259 Flicker, feces Clostridium, 795 General Salmonella, 504, 521, 526, 527 — , caecum Treponema, 1075, 1076 — , intestine Bacillus, 648, 651, 652, 653, 655, 658, 660, 662, 664, 666, 669, 671, 746, 757 Bacterium, 675, 676, 683, 685, 687, 688, 759, 760, 761, 762 Micrococcus, 269 Pseudomonas, 147, 149 Vibrio, 196 — , not pathogenic for Piscine tuberculosis Mycobacterium 883 INDEX OF SOURCES AND HABITATS 1301 Birds (continued) Turtle tuberculosis M ycobacterivm , 886 — , stomach Bacillus, 647, 648, 652, 653, 655, 658, 660, 662, 664, 666, 667, 669, 671, 746, 749, 754, 757 Bacterium, 686, 759, 760 Micrococcus, 269 Pseudomonas, 146, 147 (! rouse Salmonella, 521 ■ — , blood and intestine Spirochaeta, 1067 Guinea fowl Clostridium, 796 Hedge sparrow, stomach Micrococcus, 252, 270, 273 Parrots Miyagawanella, 1117 Salmonella, 532 — , nasal secretions Miyagawanella, 1117 Partridge Salmonella, 525 Pheasant Bacterium, 552 Clostridium , 746 Miyagawanella, 1117 Salmonella, 521 Pigeons, diseased Bacillus, 400, 652 Bacterium, 401 Erysipelothrix, 411 Haemobartonella, 1104 Mycobacterium, 881 Salmonella, 503, 532 Poultry Salmonella, 502, 505, 507, 509, 510, 511, 512, 513, 514, 516, 518, 519, 521, 522, 523, 525, 526, 528, 529, 530 — , caecum Treponema, 1075 — , erythrocytes Grahamella, 1110 — , intestine Sarcina, 292 — , nasal passages, 1294 — , not pathogenic for Bovine tuberculosis Mycobacterium, 879 — , human tuberculosis Mycobacterium, 878 — , spleen Mycobacterium, 890 — , throat Corynebacterium, 403 Quail Salmonella, 510, 521 Robin, feces Bacillus, 756 Rock dove, stomach and intestine Micrococcus, 252, 270, 276 Sarcina, 293 Snipe, intestine Chitin-digesting bacteria, 632 Sparrow, intestine Micrococcus, 270, 273, 291, 293 — , stomach Bacterium, 761 Micrococcus, 291, 293 Starling, intestine Micrococcus, 257, 263, 272, 273 — , stomach Micrococcus, 257, 263, 272, 273 Sarcina, 291 Teal, caecum Treponema, 1076 Turkey Salmonella, 510, 512, 513, 514, 519, 524, 525, 527, 528 Turkey poults Lactobacillus , 407 Salmonella, 504, 505, 514, 515, 523, 527, 528, 529 Woodpecker, intestine Micrococcus, 251, 260, 272 Sarcina, 290 Yellow hammer, intestine Micrococcus, 259, 263, 265, 272 Sarcina, 292 , stomach —Micrococcus, 259, 263, 265, 272 Birds, Diseases of Birds of prey Bovine tuberculosis Mycobacterium, 879 Canaries Infectious necrosis Pasteur ella, 554 1302 INDEX OF SOURCES AND HABITATS Birds, Diseases of (continued) Canaries (continued) Intestinal catarrh and liver changes Bacillus, 530 Septicemia Shigella, 540 Chicken, see Poultry Chicken embryos Pleuropneumonia-like disease, 1293 Cockatoo Bovine tuberculosis Mycobacterium, 879 Crossbills Infectious disease Bacillus, 661 Dove Ornithosis Miyagawanella, 1117 Ducklings Keel Salmonella, 523 Ducks Septicemia Pasteurella, 552 Pfeifferella, 554 Finches Ornithosis Miyagawanella, 1117 Fulmar petrels Ornithosis Miyagawanella, 1117 Geese Septicemia Shigella, 543 Spirochaetosis Borrelia, 1059 Cieneral Avian tuberculosis Mycobacterium, 881 Cholera-like disease Vibrio, 196 Diphtheria Bacillus, 400 Diphtheria-like disease Actinomyces, 915 Hemorrhagic septicemia Pasteurella, 549 Pleuropneumonia-like disease, 1293 Grouse Bacillus, 668 Parrot Bovine tuberculosis Mycobacterium, 879 Human tuberculosis Mycobacterium, 878 Nodules, kidney, lung, spleen Streptococcus , 342 Parrot fever Miyagawanella, 1117 Psittacosis Miyagawanella, 1117 Pheasants p43idemic in Bacterium , 552 Ornithosis Miyagawanella, 1117 Pigeon Avian tuberculosis Mycobacterium, 881 Diphtheria Bacillus, 400 Bacterium, 401 Poultry Abscesses, epidemic Sphaerophorus, 579 Chicken cholera, blood Micrococcus, 262 Chicken cholera-like disease Proteus, 489 Conjunctivitis, acute Colesiota, 1120 Coryza Coccobacillary bodies of Nelson, 1294 Diphtheria Bacterium, 674 Corynebacterium, 385 Fowl cholera Bacterium, 642 Salmonella, 521 Fowl typhoid Proteus, 489 Salmonella, 521 Hemorrhagic septicemia Pasteurella, 547, 549 Keratitis Colesiota, 1120 Lesions, tuberculous Mycobacterium, 880 Limberneck Clostridium, 780 INDEX OF SOURCES AND HABITATS 1303 Birds, Diseases of {continued) Listeriosis Listeria, 409 Ocular roup Colesiota, 1120 Ornithosis Miyaga wanella ,1117 Rhinitis, infectious Hemophilus, 589 Roup Bacteriinii, 674 Septicemia Bacterium, 642 Slipped tendon Streptococcus, 317 Spirochaetosis Spirochaeta, 1059 Tuberculosis, avian Mycobacterium, 879, 880 Tumors in abdominal cavity Actinomyces, 918 White diarrhoea Salmonella, 521 Swan Cholera Bacillus, 551 Infectious disease Bacterium, 642 Turkeys Pneumocarditis Bacillus, 662 Wild pigeons Epidemic in Bacterium, 552 Bird's Nest Myxococcus, 1042 Birds, Scientific Names Colaptes auratus, 795 Coluniba livia, 252, 270, 276, 293 Columba oenas 252, 254, 270, 271, 274 276, 292, 760, 761 Corvus cor one, 269 Ember iza citrinella, 259, 263, 265, 272, 292 Fringella carduelis, 257, 259 Fringella coelebs, 260, 266 Fulica atra, 260, 263 Gallus sp., 1075 Lagopus scoticus, 668, 1067 Larus ridibundus, 1075 Passer moritanus, 252, 270, 273, 291 293 Pious major, 251, 260, 272, 290 Pyrrhula europea, 1095 Querquedula querquedula, 1076 Squatarola squatarola, 1076 Sturnus vulgaris, 257, 263, 272, 273, 291 Blue Milk, see Dairy Products, Abnormal Milk Bovine Diseases and Sources, see Cattle Butter Actinomyces, 974 Aerobacter, 456 Bacillus, 743 Bacterium, 590, 602, 676 Chromobacierium, 234 Lactobacillus, 357 Leuconostoc, 348 Microbacferivm, 371 Micrococcus, 260, 264, 265, 266, 279 Mycobacterium, 889, 890 Nocardia, 905 Pseudomanas, 146 Sarcina, 291 Streptococcus, 325, 337, 339 Black to reddish brown discoloration Pseudomonas, 109 Fruity aroma Micrococcus, 260 May apple to strawberry odor Pseudomonas, 100, 101 Rancid Bacillus, 650, 659, 660, 814, 816 Micrococcus, 254 Skunk odor Pseudomonas, 99 Tainted Pseudomonas, 99, 100, 101, 109 Buttermilk Bacillus, 644 Propionibacterium, 374, 375, 377 Serratia, 484 Abnormal Clostridium, 822 Calf Diseases and Sources, see Cattle Cats Lungs Miyagawanella , 1118 1304 INDEX OF SOURCES AND HABITATS Cats {continued) Not pathogenic for, Avian tuberculosis Mycobacterium , 880 Human tuberculosis Mycobacterium , 878 Respiratory tract, upper Hemophilus , 586 Sputum Pastexirella, 553 Stomach Spirilhan, 218 Spirochaeta, 1069 Cats, Diseases of Actinomycosis Actinomyces, 027, 969 Bovine tuberculosis Mycobacterium , 879 Diarrhoea Salmonella, 511 Diphtheria in kittens C orynebacterium , 385 Distemper Miyagawanella , 1118 Glanders Malleomyces, 555 Hemorrhagic septicemia Pasteurella, 549 Infections Bacillus, 655 Streptococcus , 339 Influenza Miyagawanella, 1118 Miunps Spirochaeta, 1074 Nasal catarrh M i yagawan ell a, 1118 Pneumonia Brucella, 563 Pneumonitis Miyagawanella, 1118 Pseudotuberculosis Pasteurella, 550 Cattle Blood Borrelia, 1062 Clostridi%mi, 111 Micrococcus, 254 Calf vaccine lymph Bacillus, 401 C orynebacterium, 402, 406 Corneal or conjunctival discharges Colesiota, 1120 Erythrocytes Anaplasma, 1100 E per ythroz 0071, 1112 Haemobartonella, 1106 Foetus Streptococcus , 317 ' Ganglia Actinomyces, 927 Gastric mucosa Leptospira, 1078 General Salmonella, 502, 506, 509, 513, 517, 518, 521,527,529 Genital organs Brucella, 561 Glands Actinomyces, 927 Intestine Bacillus, 745, 818 Clostriditmi, 821 Micr obacterium , 370 Ri Stella, 576 Streptococcus, 322, 327 Liver Actinomyces , 926 Bacillus, 612 Recordillus, 823 Lungs Bacillus, 612 Micrococcus, 254 Lymph glands Mycobacterium , 890 Mouth cavity Actinomyces, 926 Streptococcus, 322 Mucosa, intestinal Mycobacterium, 881 Red blood cells, see Erythrocytes Runien F lav obacterium, 613 Clostridium, 820 Saliva Streptococcus, 322 Salted hides, red Pseudomonas, 110 Sarcina, 289 INDEX OF SOURCES AND HABITATS 1305 Cattle {continued) Skin Bacillus, Ibb Corynebacteriuin, 402 Mycobacterium, 890 Streptococcus, 335 Spleen Bacterium, 759 Stomach Plectridium, 823 Stomach, third Micrococcus, 272 Throat Streptococcus , 322 Udder Actinomyces, 926 Bacillus, 672 Chlorobacterium, 693 Cor?/7ie6acr?o, 203, 204, 205, 206, 207, 703 Cheese Aerobacter, 456, 457 Bacillus, 612, 647, 669, 712, 734, 743, 745, 750, 814, 815, 816, 824, 825 Bacterium, 676, 677, 678, 679, 681, 683, 686, 689, 758, 760, 761, 762, 819 Clostridium, 770, 810, 820, 822, 824 Escherichia, 452 Granulobacillus, 826 Lactobacillus, 351, 352, 356, 357, 359 Microhacterium, 370, 371 Micrococcus, 241, 243, 251, 256, 258, 264, 265, 266, 269, 270, 277, 279, 281 Propionibacterium, 373, 374, 375, 376, 377, 378, 379 5a/-ci«a, 290, 291, 292, 293 Streptococcus, 325, 326, 340, 344 INDEX OF SOURCES AND HABITATS 1307 Cheese (continued) Tetracoccus, 284 Tijrothnx, 823 Vibrio, 197 Bitter Streptococcus, 323 Black Discoloration Bacterium, 678 Gassy curd Bacillus, 672 Pasteurized Actinomyces, 968 Red Plocamobacteriuvi , 691 Rusty spot Lactobacillus, 357, 359 Cheese, Types of Blue Bacterium, 602 Brick Bacterium,. 602, 761 Brie Streptococcus , 341 Camembert Bacterium , 602 MtcrococcM^, 260, 263, 265 Sarcina, 293 Tetracoccus, 284 Cantal Streptococcus, 337 Cheddar Bacterium, 602 Pseudomonas. 697 Cream Bacillus, 664 Streptococcus, 340 Edam Chromobacterium, 233 Propionibacterium, 378 Emmental, see Swiss Gouda Bacillus, 663 Streptococcus, 342 Grana Bacillus, 756 Limburger Bacillus, 612 Bacterium, 602 Micrococcus, 261, 264 Liptauer Bacillus, 816, 826 Neuchatel Streptococcus , 342 Oka Bacterium, 602 Parmesan, see Grana Bacillus, 737, 756 Richelieu Micrococcus, 695 Schlosskase Streptococcus, 342 Soft Bacillus, 649 Salmonella, 529 Swiss Bacillus, 648, 657, 737, 739, 752 Bacterium, 688, 759, 760 M7c;-ococci/s, 255, 256, 257, 259, 261, 263, 264, 269, 270, 273, 275, 277, 281 Propionibacterium, 373, 374, 379 Sarcina, 290, 291 Streptococcus, 323, 338 Tilsit Plocamobacterium , 691 Propionibacterium , 378 Chitin Digesting Bacteria Bacillus, 632 Bacterium, 632 Concrete, see Mineral Sources Condensed Milk, see Dairy Products Contamination Bacillus, 712, 717, 748 Bacterium, 604, 673, 682 Cellulomonas, 622 Chromobacterium, 232 Clostridimn, 798 Corynebacterium, 401 Nocardia, 905, 915 Oospora, 976 Planococcus, 281 Pseudomonas, 174 Staphylococcus, 282 Tuberculin Mask Nocardia, 905 ^^accine Oospora, 976 Cream, see Milk and Cream Cutting Compound, see Mineral Sources 1308 INDEX OF SOURCES AND HABITATS Dairy Products, also see Butter, Cheese, and Milk and Cream Abnormal milk Bitter flavor Bacillus, 648 Micrococcus, 265 Blue milk Pseudomonas, 93, 145 Fetid odor Viscobacterium, 691 General Achrombacler , 609 Bed milk Bacterium,, 601 Micrococcus, 256 Ropy milk, also see Slimj- milk Achromobacter, 427 Alcaligenes , 414 Lactobacillus, 695 Micrococcus, 258, 268, 271 Streptococcus, 344, 702 Slimy milk, also see Ropy milk Achromobacter , 425 Bacillus, 654, 746 Bacterium, 604, 675 Clostriditim, 789 Flavobacterium, 434, 442 Micrococcus, 268, 271 Soapy milk Bacillus, 660, 667, 758 Pseudon}onas, 145, 149, 150 Tainted Bacillus, 660 Aroma bacteria Streptococcus, 339 Caseinate solution Achromobacter, 692 Condensed milk Leuconostoc, 348 Thermobacterium , 364 Evaporated milk Bacillus, 713, 738 Pseudomonas, 103 Fermented milk Dadhi Streptothrix, 364 General Sarcina, 292 Gioddu Bacillus, 362 Kefir Bacillus, 746 Lactobacillus, 351, 357, 359 Streptococcus , 338 Kumys Bacillus, 362 Leben Streptobacillus , 364 Matzoon Flavobacterium, 441 Sarcina, 252 Mazun Bacillus, 748 Bacterium, 362 Piima Streptococcus, 342 Taette Lactobacillus , 695 Streptococcus, 702 Taette, false Bacterium, 675 Yoghurt Lactobacillus, 355, 362, 364 General Bacillus, 732, 733 Mrcrococc?/^, 237, 239, 240, 241 Mycobacterium, 889 Propionibacterium, 373, 374, 375, 376, 377, 378, 379 Pseudomonas, 100 Ice cream Bacillus, 612, 670 Malted milk Bacillus, 756 Milk powder Bacillus, 733, 755, 756 Milk powder (spray process) Streptococcus , 328 Sour milk Bacillus, 756 Lactobacillus , 352, 362, 364 Starters Leuconostoc, 347, 348 Streptococcus, 325 Whey Bacillus, 755, 756 Bacterium, 687 Dairy Utensils Alcaligenes, 414 Bacillus, 732, 733 Microbacteriuvi, 370, 371 INDEX OF SOURCES AND HABITATS 1309 Dairy Utensils {continued) Micrococcus, 238, 239, 24U, 241 Pseudomonas, 100 Creamery equipment Pseudomonas, 99 Farm utensils Sarcina, 294 Filter cloth Bacillus, 735 Milk can Serratia, 481 Milking machine Sarcina, 294 Dairy Wastes Achromobacter, 628 Flavobacteriurn, 433 Pseudomonas, 177, 178 Decomposing Materials Agar-digesting bacteria, see Agar Di- gesting Bacteria Cellulose-digesting bacteria, see Cell- ulose Digesting Bacteria ' Cellulose fibers Sorangium, 1024 Chitin-digesting bacteria, 632 Composts Cellulomonas, 618 Micromonospora, 980 Strepiomyces, 957 Thiobacillus, 79, 81 Crab shells Bacterium, 632, 633 Farinaceous materials Clostridium, 824 General Bacillus, 711, 715, 730, 733, 734, 735, 741, 743 Bacterium, 682 Myxococcus, 1042 Spirillum, 701 Grass Actinomyces, 973 Leaves Bacillus, 756 Myxococcus, 1042 Lichens Archangium, 1018, 1019 Myxococcus, 1042 Litmus solution Bacillus, 743, 751 Marine vegetation Cytophaga, 1014, 1015 Organic Matter Chondrococcus, 1046 Cytophaga, 1014 Polyangium, 1028, 1029, 1030 Sorangium, 1023 Fzfcrro, 201, 203 Paper Myxococcus, 1042 Plant residues Bacterium, 824, 825 Clostridium , 825 Cytophaga, 1013 Starchy materials Bacillus, 722 Sugar refinery waste Lampropedia, 844 ^'egetable Bacillus, 721 Spirillum, 214 Vegetation Bacillus, 815, 819 Thiothrix, 989 Wood Chondromtjces, 1037, 1039 Clostridium, 825 Myxococcus, 1042 Synangium , 1033 Watermelon Bacillus, 748, 751 Water plants Lampropedia, 844 Dogs Bile Bacter aides, 566 Blood Leptospira, 1078 Spirochaeta 1066 Streptococcus, 343 Conjunctiva Corynebacierium , 406 Erythrocytes Haemobartonella, 1104 Fatal illness Rickettsia, 1096 Feces Bacillus, 818 Salmonella, 514, 529, 530 1310 INDEX OF SOURCES AND HABITATS Dogs (continued) Genital organs Salmonella, 514, 529, 530 Intestine Bacterium, 680 Salmonella, 514, 529, 530 Spirochaeta, 1068 Liver Acuformis, 812 Nasal mucosa Zuberella, 578 Nasal mucus Bacillus, 740 Not pathogenic for Mycobacterium , 880 Nocardia, 896 Preputial secretions Hemophilus, 588 Red blood cells, see Erythrocytes Skin Corynethrix, 406 Stomach Bacillus, 651, 671 Spirillum, 217, 218 Spirochaeta, 1069 Throat Streptococcus , 345 Urine Bacillus, 740 Dogs, Diseases of Abort ion Brucella, 562 Abscess Micrococcus, 273 Actinomycosis Actinomyces, 927 Bronchitis Nocardia, 919 Cancerous ulcers Spirochaeta, 1074 C'atarrh Pseudomonas, 146 Cerebromeningitis Nocardia, 919 Distemper Brucella, 563 Endometritis Bacterium, 677 Glanders Malleomyces, 555 Haemobartonellosis Haemobartonella, 1102 Hemorrhagic septicemia Pasteurella, 549, 550, 553 Human tuberculosis Mycobacterium, 878 Leukemia Bacterium, 680 Streptococcus, 337 Lymphadenitis Nocardia, 919 Measles Streptococcus , 341 Peritonitis Actinomyces , 915 Phlegmon Actinomyces , 915 Plague Spirochaeta, 1068 Pleuropneumonia-like disease Aster ococcus, 1292 Pneimionia Brucella, 563 Purulent infectious, urinary tract C orynebacterium , 389 Pyorrhoea Leptothrix, 366 Rabies, vaccine contaminant Flavobacterium, 437 Rabies-like disease Bacillus, 663 Ringworm Actinomyces, 916 Uremia Leptospira, 1078 Dung, see Manure (Dung) Dust, also see Air. Achromobacter, 424, 425, 427 Actinomyces, 968, 970 Bacillus, 712, 717, 723, 725, 726, 729, 730, 731, 732, 733, 734, 735, 736, 737, 738, 739 Bacterium, 605, 682 Chromobacterium, 233 Clostridium, 786, 817, 820 Diplococcus, 309 .1/icrococcws, 237, 239, 240, 243, 244, 253, 255, 256, 257, 258, 259, 260, 262, 264, 265, 269, 273, 275, 276, 277, 278, 279, 280, 281 INDEX OF SOUKCES AND HABITATS 1311 Dust (continued) Miyagawanella, 1117 Mycobacterium, 889, 890 Sarcina, 286, 290, 292, 293 Serratia, 484 Streptococcus, 316, 340 Streptomyces, 935, 936, 949 Dusts Stable Micrococcus, 265 Street Clostridium, 786, 820 Eggs, see Foods and Foodstuffs Evaporated milk, see Dairy Products Feces, Animal, see Manure (Dung) Feces, Himian Also see Human Sources, Intestine Aerobacter, 456, 457 Alcaligenes, 413, 416 Bacillus, 612, 647, 651, 652, 654, 656, 661, 662, 665, 666, 668, 721, 736, 737, 738, 744, 746, 748, 749, 753, 755, 756, 813, 817, 818, 825, 826 Bacterium, 601, 607, 673, 675, 681, 683, 687, 759, 760, 761, 762 Bacter aides, 567, 568, 569, 570, 571, 572, 573, 574, 575 Bifidobacterium, 369 Catenabacterium, 368 Clostridium, 773, 774, 782, 785, 787, 788, 791, 793, 794, 796, 797, 799, 800, 801, 802, 809, 810, 811, 812, 820, 821, 822, 824, 826 Corynebacterium, 401 Eberthella, 533, 534 Escherichia, 447, 450, 451, 452, 453 Eubacterium, 368 Granulobacillus , 826 Kurthia, 613 Lactobacillus, 353, 354, 357, 359, 362 Microbacterium, 370 Micrococcus, 252, 253, 257, 259, 260, 265, 266, 270, 272, 274, 275, 276, 277, 278, 279, 280, 281, 695 Microspira, 202 Proteus, 488, 490, 491 Pseudomonas, 90, 148 Ramibacterium 369 Salmonella, 502, 504, 505, 506, 510, 511, 512, 513, 514, 515, 517, 518, 519, 52o[ 522, 524, 525, 526, 527, 528, 529, 530, 531 Shigella, 537, 538, 539, 540, 542, 543, 544 Spirochaeta, 1067, 1069, 1070 Staphylococcus, 282 Streptococcus, 322, 323, 326, 327, 330 333, 335, 339, 341, 343 Vibrio, 204, 205 Feces, Infant Micrococcus, 257 Neisseria, 301 Pseudomonas, 198 Staphylococcus, 281 Fermenting and Fermented Materials Agave americana, sap Pseudomonas, 106 Alcohol infusions Bacterium, 675 Ale, bottled Acetobacter, 692 Beer, see Beer Beet juice Acetobacter, 189 Beets Bacillus, 656, 664, 752, 757 Bacterium, 675, 678, 686, 761 Lactobacillus, 357 Beverages Acetobacter, 181, 182 Bread dough Bacillus, 657, 664, 742 Bacterium, 683 Lactobacillus, 357, 359, 361, 363 Cabbage, see Sauerkraut Cereal mash Lactobacillus, 363 Corn mash Clostridiuyn, 781, 825 Lactobacillus, 357 Dates Acetobacter, 184 Dough Bacillus, 742 Bacterium, 683 Ensilage, see Silage. Fermented milk drinks, see Dairy Products, Fermented milk 1312 INDEX OF SOURCES AND ttABlTATS Fermenting and Fermented Materials (continued) Figs Acetobacter , 184 Flax retting, see Retting, Flax Fodder Bacillus, 750, 755 Fruits Acetobacter, 181, 182, 183, 184 Ginger beer Bacterium, 362 Grain mash Acetobacter, 182 Bacillus, 824 Granulobacter, 822, 824 Lactobacilhis , 356, 357, 359, 363 Hay Bacillus, 740 Hemp retting, see Retting, Hemp Hydrogen peroxide solutions Acetobacter, 189 Infusions Bacillus, 672 Kenaf (Hibiscus) retting see Retting, Kenaf Kombucha from tea infusions Acetobacter, 189 Malt Bacterium, 677, 678, 679, 688, 760, 761 Malt beverages Acetobacter, 182 Malt infusion Micrococcus, 263 Malt mash Sarcina, 287 Mash, dried persimmon Acetobacter, 184 Mash, spoiled Pediococcu^, 249 Mash, vegetable Lactobacillus, 356, 357, 359, 361 Methane fermentation in swamps Sarcina, 287 Milk, Fermented see Dairy Products, Fermented Milk Molasses Clostridium, 781 Lactobacillus, 359, 363 Pickles Bacillus, 648 — , cauliflower Lactobacillus, 357 — , cucumber Lactobacillus, 357 — , tomato Lactobacillus, 357, 359 Plant juices Pseudo?ndnas , 106 Plant materials Clostridium, 772 Lactobacillus, 347 Streptococcus, 325, 326, 327, 340 Potato mash Amylobacter, 813 Clostridium, 781, 808, 819 Lactobacillus, 356, 357, 361 Pulque Pseudomonas, 106 Streptococcus, 338 Sake Lactobacillus, 363 Sauerkraut Bacillus, 657, 750 Bacterium, 675 Lactobacillus, 357, 359 Pseudomonas, 146 Silage Bacterium, 602 Clostridium, 785 Lactobacillus, 359 Propionibacterium, 376 Streptococcus, 336 Slimy fermentation Bacillus, 749 Soybean cake Bacillus, 757 Tobacco Bacterium, 674, 682, 685 Urea Staphylococcus, 282 Urine ymcrococci^, 238, 250, 260, 266, 267, 269, 279, 280 Sarcina, 289, 293 Staphylococcris , 282 Vegetables Acetobacter, 181, 182 Lactobacillus, 347, 356, 357, 359, 361 Vinegar ^ceio&ac^er, 181, 182, 183, 186, 187, 188 INDEX OF SOURCES AND HABITATS 1313 Fermented Materials (continued) Vinegar (continued) Bacillus, 756 Bacterium, 682 — , quick process Acetobacter, 187, 188 — , wine Acetobacter , 187, 188 Wine, see Wine Yeast, see Yeast Fishes African Mudfish, bluod Spirochaeta, 1067 Anchovy pickle Pediococcus, 250 Anchovy, salted Vibrio, 204 Blenny, intestine Treponema, 1045 Brochet Bartwiella, 1108 Bullhead, Marine Treponema, 1074 Carp, pathogenic for Bacterium, 642 Proteus, 491 Pseudomonas, 102, 149 Mycobacterium, 883, 884 Catfish Serratia, 462 Codfish — , feces Achroynobacter, 420, 423, 426, 427 Flavobacteriun, 434 — , intestine Achromobacter , 425 Shigella, 544 — , red salted Micrococcus, 259 Pseudomonas, 110 Sarcina, 289 — , slime Achromobacter, 420, 423, 426, 427 Flavobacterium, 434, 436 Croaker Mycobacterium, 884 Dogfish Flavobacterium, 436 — , slime and feces Achromobacter, 420 Eels Bacterium, 673 Mycobacterium, 883 Vibrio, 208 General Bacillus, 753 Pseudomonas, 102, 109, 149 — , blood Rickettsia, 1097 Spirochaeta, 1068 — , intestine Chitin-digesting bacteria, 632 — , rectum Treponema, 1076 — , skin Bacterium, 606, 612 Micrococcus, 246 Haddock Spirochaeta, 1065 — , slime Flavobacterium, 438, 440, 441 Hake Flavobacterium, 439 Halibut Pseudomonas, 145 — , skin Flavobacterium, 429, 434, 437 Herring, salted Sarcina, 292 Kilifish Pseudomonas, 109 Lamprey eel Bartonella, 1108 Marine fishes Mycobacterium, 883 Pseudomonas, 147 Spirochaeta, 1067 Treponema, 1075 Perch, erythrocytes Grahamella, 1110 Pollack Spirochaeta 1069 Salmon, Blue-black Chondrococcus, 1047 Salmon eggs Achromobacter, 425, 692 , diseased Pseudomonas, 700 Salted fishes Micrococcus, 259, 266, 268, 292 Pseudomonas, 110 Sarcina, 289 Vibrio, 2(h 1314 INDEX OF SOUECES AND HABITATS Fishes (continued) Sardines, salted Vibrio, 204 — , stomach Euhacierium, 367 Sea bass Mycobacterium, 884 Sergeant major Mycobacterium, 884 Shark, blood Borrelia, 1064 Skate, slime and feces Flavobacterium, 434, 439 Tench Bartonella, 1108 Trout Bacterium, 686 Whiting Microspironema, 1064 Fishes, Diseases of Carp — , tumors Mycobacterium, 883, 884 — , red spots Bacterium, 642 Eels, diseased Bacterium, 673 Mycobacterium, 883 Vibrio, 203 General — , epidemic infection of fishes Bacillus, 751 Vibrio, 197 — , fresh water fishes Hemorrhagic septicemia Pseudomonas, 102, 103 — , skin lesions Pseudomonas, 102 — , necrotic ulcers Rickettsia, 1097 — , necrosis of the liver Chondrococcus, 1047 Mycobacterium, 884 Kilifish, skin lesions Pseudomonas, 109 Marine fishes Pseudomonas, 149 , infected skin and muscle Pseudomonas, 109 , tuberculosis Mycobacterium, 884 Trout, furunculosis Bacterium., 686 Salmon eggs Achromobacter, 692 Pseudomonas, 700 Fishes, Scientific Names Abudefduf mauritii, 884 Acropoma japonicum, 636 Ameiuras melas, 462 Blennius pavo, 1075 Box boops, 1073 Centropristes striatus, 884 Clarias angolensis, 1067 Coelorhynchus sp., 636 Cottus bubalis, 1074 Cyprinus carpio, 491, 642, 884 Cyprinus sp., 102, 149 Esox lucius, 1108 Fundulus parvipinnes, 109 GadMS callarias, 420, 423, 425, 426, 427, 434, 436, 544 Gadus minutus, 1067 Gadus pollachius, 1069 Hippoglossus hippoglossus, 429, 434, 436, 437 Lepadogaster bimaculatus, 1075 Melanogrammus aeglefinus, 438, 440, 441 Merlangus merlangus, 1064 Micropogon undulatus, 884 Oncorrhynchus nerka 1047 Pelamys sarda, 1068 Perca fluviatilis, 1110 Petromyzon marinus, 1108 Physiculus japonicus, 636 /^aja crinacea, 434, 439 Saccobranchus fossilis, 753 Squalus acanthias, 420, 427, 436 Tetraodon fahaka, 1097 Tinea tinea, 1108 Trachurus japonicus, 694 Trigla lucena, 1076 Urophycis tenuis, 439 Fomites Hairbrush Micrococcus, 253, 275 Hospital shirt Serratia, 484 INDEX OF SOURCES AND HABITATS 1315 Foods and Foodstuffs Anchovies, pickled Micrococcus, 250 Asparagus, boiled Bacillus, 742, 748 Bacterium, 759 Bacon, tainted Vibrio, 702 Beans, salted, purple discoloration Pseudomonas, 109 Beef extract Bacillus, 734 Beet juice, sugar Leuconostoc, 348 Blood sausage Clostridium, 788 Blutwurst Clostridium , 788 Bread Bacillus, 661 Bacterium, 680, 759 — , rye Bacillus, 711, 750 — , slimy Bacilbis, 711, 758 Bacterium, 760 Butter, see Butter Candy Bacillus, 756 Canned beans Bacillus, 750 — , beets, blackened Bacillus, 739 — , blueberries Bacillus, 735 — , carrots Bacillus, 742 — , corn, spoiled Bacillus, 734, 756 — , corn, sulfur stinker spoilage Clostridium, 803 — , evaporated milk Bacillus, 713, 738 Pseudomonas, 103 — , foods Bacillus, 713, 730, 731, 734, 756 Clostridium, 779, 797 — , goods, spoiled Bacillus, 730, 731, 734 — , goods, spoiled, non-acid Clostridium, 785, 803 — , macaroni, spoiled Bacillus, 317 — , peas Bacillus, 737, 756 — , pimipkin Bacterium, 609 — , pumpkin, swells Bacillus, 730, 731 — , salmon, spoiled Bacillus, 817 — , sardines Ser ratio, 483 — , spinach Bacillus, 825 — , string beans, spoiled Bacillus, 734, 748 — , tomatoes Bacillus, 756 — , vegetables, flat sours Bacillus, 734 Catsup Lactobacillus, 359 Cheese, see Cheese Codfish, reddened salt Bacillus, 667, 742 Flavobacterium , 442 Pseudomonas, 110 Corn meal Bacterium, 679 Crab meat, musty odor Achromobacter, 425 Cream, see Milk and Cream Dairy Products, see Dairy Products Dates, commercially packed Bacillus, 756 Eggs, black rot Proteus, 490 — , cooked Bacterium, 761 — , duck Salmonella, 517 — , hen's Bacillus, 653, 654, 657, 659, 663, 672, 747, 750 Pseudomonas, 147, 148, 149, 150, 179 — , musty Achromobacter, 425 Pseudomonas, 148, 179 — , powdered Salmonella, 510, 512, 513 1316 INDEX OF SOURCES AND HABITATS Foods and Foodstuffs (continued) Fish conserves ' Clostridium, 821 — , herring, rusty Pseudomonas, 110 — , salted Bacillus, 658 Ristella, 576 ■ — , semi -dried Flavobacterium , 69-4 Food conserves Bacillus, 741 Gelatin, spoiled Bacillus, 756 General Serratia, 481, 482 Grapes, Spanish dried Bacterium, 624 Ham, salted Euhacterium, 367 — , sour Clostridium, 784 Horseradish, ground Bacillus, Ibl IMargarine Bacillus, 662, 667 Bacterium, 681, 685, 689 Meat Bacterium, 678 Micrococcus, 258, 272, 281 Streptococcus, 338 - — , extract Bacillus, 648, 651, 740, 745, 746, 749 Bacterium, 678, 760 Micrococcus, 255, 267 Streptococcus, 339 — ■, pies Salmonella, 531 — , spoiled Bacillus, 749 Milk, see Milk and Cream Mincemeat, canned Bacillus, 757 Oranges Butylobacter , 825 Oysters Inflabilif, 823 Pickles Bacillus, 648, 754 Salmonella, 519 Plum preserves Bacillus, 753 Pork Clostridium, 802 Potato, cooked 5acz7Z7is, 664, 741,748 Rice, cooked in chicken broth Serratia, 484 Salad dressing Lactobacillus, 363 Sausage Bacillus, 749, 751, 752, 816 Clostridium, 778, 779 Salmonella, 530 Sugar, also see Sugar Bacillus, 742, 745, 747 Leuconostoc, 347, 348 Micrococcus, 260 Spirillum, 217 — , beet Bacilltis, 747 — , factories, frog spawn fungus Leuconostoc, 347, 348 Spirillum, 217 Tomato juice Bacillus, 713 Tomato products, spoiled Lactobacillus, 357, 359 Truffles, cooked Bacillus, 757 Wiener skins Tetracoccus, 284 Wurst Micrococcus, 258, 272, 281 Goats Cerebrospinal fluid Streptococcus, 338 Corneal or conjunctival discharges Colesiota, 1120 General Salmonella, 506 Goats, Diseases of Abortion Brucella, 561 Agalactia Anulojnyces , 1292 Glanders Malleomyces, 555 INDEX OF SOURCES AND HABITATS 1317 Goats, Diseases of {continued) Heartwater Cowdria, 1094 Hemorrha,gic septicemia Pasteurella, 549 Keratitis Colesiota, 1119 Lesions Nocardia, 900 Ophthalmia, infectious Colesiota, 1120 Pleuropneumonia, bovine Asterococcus, 1292 Tuberculosis, bovine Mycobacteriu7n, 879 Guinea Pigs Blood Rickettsia, 1086, 1088, 1089 Spirochaeta, 1066, 1070 Spiroschaiidinnia, 1071 Spironema, 1070 Cadaver BacillvjS, 815 Caecum Metahacterium , 762 Treponema, 1075 Erythrocytes Haemobartonella, 1104, 1108 Erythrocytes, Peruvian guinea pigs Haemobartonella, 1106 Inoculated with soil Bacillus, 660, 817, 825, 826 Hiblerillus, 822 Intestine Cristispirilla , 1057 Heliconema, 1064 Oscillospira, 1004 Sarcina, 287 Intestine and genital organs Salmonella, 506, 527 Liver Spirochaeta, 1065 Lymph glands BacilltLS, 659 Monocytes Ehrlichia, 1095 Mucus, intestinal Bacillus, 651 Nasal mucosa Zuberella, 578 Normal animals Veillonella, 304 Not pathogenic for Johne's disease Mycobacterium , 881 Red blood cells, see Erythrocytes Tuberculosis, avian Mycobacterium, 880 Tuberculosis, piscine Mycobacterium, 883, 884 Tuberculosis, snake Mycobacterium, 885, 886 Tuberculosis, turtle Mycobacterium, 886 Stomach Klebsiella, 459 Tunica vaginalis Rickettsia, 1086, 1089 Guinea Pigs, Diseases of Anthrax Bacillus, 720 Brucellosis Brucella, 561, 562, 563 Cervical adenitis Bacteroides, 575 Epizootic Bacterium, 681 Glanders Malleomyces, 555, 556 Hemorrhagic septicemia Pasteurella, 549, 550, 551, 553, 554 Infections Gaffkya, 284 Nocardia, 913 Listerosis Listeria, 409 Lymphadenitis Streptococcus, 317 Maculatum disease Rickettsia, 1098 ^Melioidosis Malleomyces , 556 Septicemia Pseudomonas, 146 Skin abscesses Neisseria, 301 Tuberculosis, bovine Mycobacterium, 878 Tuberculosis, human Mycobacterium, 878, 879 Tuberculosis-like disease Pasteurella, 553 Tuberculosis, pulmonary Bacillus, 651 1318 INDEX OF SOURCES AND HABITATS Hail Bacillus, 744 Micrococcus , 269, 279 Hogs General Salmonella, 502, 504, 505, 507, 509, 510, 513, 514, 519, 521, 522, 525, 526, 527, 530 Genital organs Brucella, 562 Intestine Bacillus, 753 Lymph glands Salmonella, 505, 510, 513, 514, 518, 524, 528, 529 Liver, necrotic foci Sphaerophorus, 579 Ovary Bacillus, 757 Peritoneal fluid Bacillus, 666 Hogs, Diseases of Abortion Brucella, 561, 562 Abscesses Corynebacterium, 388 Vibrio, 206 Actinomycosis Actinomyces, 926 Anthrax Bacillus, 720 Blood in hog cholera Borrelia, 1063 Bronchopneumonia Bacillus, 657 Calcareous deposits in muscles Actinomyces, 972 Caseous suppuration C orynebacterium , 406 Conjunctivitis Rickettsia, 1120 Cutaneous lesions Spirochaeta, 1069 Diarrhoea Bacillus, 826 Erysipelas Erysipelothrix, 411 Heartwater-like disease Rickettsia, 1097 Hemorrhagic septicemia Pasteurella, 548, 549 Hog cholera Bmrelia, 1063 Salmonella, 509, 531 Infections Corynebacterium, 391 Influenza Hemophilus, 586 Listeriosis Listeria, 409 Marseille's disease Bacillus, 662 Measles Streptococcus, 341 Pyorrhoea Leptothrix, 366 Septicemia Streptococcus, 317 Swine erysipelas Bacillus, 652 Erysipelothrix, 411 Swine fever Salmonella, 509 Swine plague (Hog cholera) Micrococcus, 278 Tuberculosis, avian Mycobacterium, 880, 881 Tuberculosis, bovine Mycobacterium, 879 Ulcers., intestinal Borrelia, 1063 Horses Blood Nocardia, 897 Spirochaeta, 1066 Female genital tract Klebsiella, 459 Streptococcus, 340 Foetus Streptococcus, 317 General Rickettsia, 1097 Salmonella, 518, 529 Intestine Bacillus, 694 Bifidobacterium, 369 Streptococcus, 323, 327 Zuberella, 577 Hock- joint (foals) Nocardia, 910 INDEX OF SOURCES AND HABITATS 1319 Horses (continued) Large intestine Bacillus, 612, 747, 749, 755, 813, 814 815, 817, 818, 826, 827 Clostridium, 783, 799 Euhacterium, 368 Gaffkya, 284 Hiblerillus, 822 Inflahilis, 823 Micrococcus, 261 Liver Malleomyces, 555 Sabnonella, 507 Nasal passages Corynebacterium, 385 Xot pathogenic for Avian tuberculosis Mijcohacteriujn^ 880 Pneumonia Corynethrix, 407 Respiratory tract Streptococcus, 318, 337 Saliva Nocardia, 975 Slcin Corynethrix, 406 Spleen Malleomyces, 555 Tliroat Streptococcus, 345 Urine Bacterium, 642 Pediococcus, 250 Sarcina, 291 Horses, Diseases of Abortion Brucella, 561, 562 Salmonella. 506 Streptococcus, 336 Abscesses on jaws Nocardia, 920 Acne pustules Bacillus, 658 Corynebacterium, 401 Botryomycosis Micrococcus, 253 Endometritis Streptococcus, 317 Gangrenous dermatitis Sphaerophorus, 579 General ■ Streptococcus, 341 Glanders Malleomyces, 551, 555 Hemorrhagic septicemia Pasteurella, 549, 550, 551, 553 Infections, genitourinary system Klebsiella, 459 Influenza Streptococcus, 340 Joint ill of foals Shigella, 542 Lymphangitis, ulcerative Corynebacterium, 389 Nasal secretion in glanders Bacterium, 683 Pneumonia Bacterium, 684 Streptococcus, 339 — , infectious of foals Corynebacterium, 391 Purulent infections, urinary tract Corynebacterium, 389 Pus, respiratory tract Streptococcus, 318 Ringworm Actinomyces, 916 Stomatitis Treponema, 1074 Strangles Streptococcus, 318 Ulcerative lesions Corynebacterium, 380 Wounds Corynebacterium, 385 Human Diseases Abscesses Alcaligenes, 413 Micrococcus, 242, 244 Proteus, 488, 490 Pseudomanas, 89 Sabnonella, 507 Streptococcus, 333 Veillonella, 303 — , abdominal Bacillus, 815 — , alveolar Bacterium, 678 — , brain Bacillus, 656, 666, 815 Capsularis, 577 Nocardia, 897 1320 INDEX OF SOURCES AND HABITATS Human Diseases {continued) Abscesses, cervical Actinomyces, 916 — , chest Nocardia, 920 — , dental Aerobacter, 456 Bacillus, 650 — , ear Corynebacterium, 402, 403 — , facial Bacterium, 607 — , foot Actinomyces, 973 — , iliac Oospora, 922 — , inguinal Nocardia, 976 — , intestinal Bacillus, 816 — , jaw Actinomyces, 917 Nocardia, 921 Proactinomyces , 923 Streptothrix, 924 — , kidney Clostridiian, 825 — , liver Bacillus, 400, 660, 666 Bacterium, 677 Bacteroides, 567 C ohnistreptothrix , 975 Proteus, 490 Sphaerophorus, 579 Fibrzo, 205 — , lung Bacillus, 667 Ristella, 575 Sphaerophorus, 579 — , mouth Corynebacterium, 402 — , multiple Nocardia, 921 — , osseus Diplococcus, 310 — , palm Actinomyces, 971 — , parotid Streptomyces , 963 — , perianal Sphaerophorus, 580 — , periuterine Veillonella, 303 — , pulmonary Actinomyces, 917 — , rectal Eberthella, 534 — , skin Streptococcus, 333 — , subcutaneous Actinomyces, 916 — , teeth Actinomyces , 917 ^ero6ac1 albus (Galactococcus) , 250 albus (Micrococcus), 242, 249, 251, 252, 253, 257, 258, 260, 261, 262, 263, 264, 265, 266, 267, 268, 269, 272, 273, 274, 275, 276, 277, 279, 280, 281, 347, 891 albus I (Micrococcus) , 257 albus II (Micrococcus) , 252, 270 albus (Pediococcus), 249 albus (Proactinomyces) , 923 albus (Staphylococcus), 242, 1140, 1141, 1142 albus (Streptococcus), 337 albus (Streptomyces), 933, 943, 947, 949 albus var. a (Actinomyces) , 934 alb^ls var. acidus (Actinomyces) , 915, 934 albus var. cretaceus (Actinomyces) , 934 albus var. maltigenes (Micrococcus) , 695 albus var. ochraleuceus (Actinomyces), 934, 968 albus var. tossica (Actinomyces), 918 aZ6MS var. toxica (Actinomyces) , 934 aibiis acidus (Actinomyces) , 915, 934 aZbus anaeruhiescens (Bacillus), 648 aZfews asporogenes (Actinomyces) , 968 aZ&us cadaveris (Bacillus), 669 aZ6ws cadaveris (Bacterium), 669 albus jluidificans (Micrococcus), 257 albus liquefaciens (Micrococcus) , 274 albus liquefaciens (Staphylococcus) , 281 aZftus non liquefaciens (Coccus), 261 aibus non liquefaciens (Staphylococcus), 281 aZbws putidus (Bacillus), 648 aZ6us putridus (Bacillus) , 648 aibws urinae (Micrococcus) , 279 albus -vulgaris (Actinomyces), 968 alcalescens (Bacillus), 450 alcalescens (Escherichia), 450 alcalescens (Micrococcus) , 303 alcalescens (Veillonella), 303 alcalescens var. gingivalis (Veillonella), 304 alcalescens var. minutissima (Veillonella), 304 alcalescens var. syzygios (Veillonella), 304 alcaliaromaticum (Achromobacter) , 416 alcaliaromaticum (Bacterium), 416 alcalifaciens (Bacillus), 400 alcalifaciens (Eberthella) , 533 Alcaligenes, 10, 21, 30, 32, 412, 416 alcaligenes (Acuformis) , 801 alcaligenes (Bacillus), 413 alcaligenes (Bacterium) , 413 alcaligenes (Clostridium), 8M, alcaligenes (Palmula), 801 1394 INDEX OF NAMES OF GENERA AND SPECIES alca-amet alcaligenes (Vibrio), 413 alcalinofoetidus (Alcaligenes) , 416 alcalinofoetidus (Bacillus), 416 alcalinus (Thermobacillus) , 734 alcalophilus (Bacillus), 738 aids (Klebsiella), 459 alcoholphilus (Lactobacillus) , 363 aleuritidis (Bacterium), 131 aleuritidis (Phytomonas) , 131 aleuritidis (Pseudomonas), 131 alfalfae (Bacterium), 165 alfalfae (Phytomonas), 165 alfalfae (Pseudomonas), 165 alfalfae (Xanthomonas), 165 algeriense (Bacterium), 673 alginicum (Bacterium), 641 alginovorum (Bacterium), 626 algosus (Vibrio), 702 aliphaticum (Bacterium), 673 aliphaticum liquefaciens (Bacterium), 673 alkalescens (Bacillus), 539 alkalescens (Bacterium), 539 alkalescens (Eberthella) , 539 alkalescens (Proshigella) , 539 alkalescens (Shigella), 539, 540, 543 allantoicus (Streptococcus) , 337 allantoides (Bacillus), 673 allantoides (Bacterium) , 673 aWer (Leuconostoc) , 346 alliariae (Bacillus), 477 alliariae (Erwinia), 477 a/iiz (Bacillus), 697 aWu' (Pseudomonas) , 145, 697 alliicola (Phytomonas) , 136 alliicola (Pseudomonas), 136 allium (Bacterium) , 145 alluvialum (Mycobacterium) , 919 alma (Cellulomonas), 620 almquisti (Actinomyces), 934, 947, 968 almus (Bacillus), 620 oinz (Actinomyces) , 968 aZoes (Bacterium), 758 alopecuri (Bacillus), 738 aZp/ia (Bacillus), 648, 738 aip/ia (Micrococcus) , 279 aip/ia (Oospora), 934 alpha (Phagus), 1141 aZp/io (Streptothrix) , 976 alpha (Tarpeia), 1268 alpinus (Bacillus), 738 altendorf (Salmonella), 506 alternans (Zygoplagia), 13 alutacea (Pseudomonas), 178 alutacea (Sarcina), 290 alutaceum (Bacterium), 673 alvearis (Cryptococcus) , 337 alvearis (Streptococcus), 337 alvei (Bacillus), 723, 724, 728, 745 alveicola (Proteus), 490 alveolaris (Bacillus), 738 aZri (Micrococcus) , 274 amabilis (Bacillus), 648 amabilis (Bacterium), 648 amager (Salmonella) , 524 amaracrylus (Aerobacillus) , 720 amaracrylus (Bacillus), 720 amaranthi (Bacterium), 178 amaranthi (Phytomonas), 178 amaranti (Pseudomonas) , 178 amarifaciens (Micrococcus), 238 amarijicans (Bacillus), 716 . amarillae (Bacillus), 648 amarillum (Plectridium) , 814 amarus (Bacillus), 648, 738 ambigua (Eberthella), 536 ambigua (Pseudomonas), 103, 697 ambigua (Shigella), 636 ambigua (Spirochaeta) , 1065 ambigua (Treponema) , 1065 ambiguum (Achromobacter) , 103 ambiguum (Bacterium) , 536, 673, 697 ambiguus (Bacillus), 103, 536 ambratus (Streptococcus), 337 americana (Cohnistreptothrix) , 974 americanum (Clostridium), 819 americanus (Actinomyces), 975 americanus (Nitrosococcus) , 71 americanus (Proteus), 490 amerimnus (Bacillus), 648 amersfoort (Salmonella), 511 amethystina (Pseudomonas) , 233 amethystinum (Chromobacterium) , 232 amethystinus (Bacillus), 232, 233 amethystinus (Bacterium), 232 1395 amet-anae INDEX OF NAMES OF GENEEA AND SPECIES amethystinus mobilis (Bacillus) , 233 amethystinus mobilis (Bacterium) , 233 amforeti (Bacterium) , 673 amherstiana (Salmonella), 515 aminovorans (Bacillus), 738 ammoniae (Proteus), 490 ammoniae (Salmonella), 490 ammoniagenes (Alcaligenes) , 416, 607 ammoniagenes (Bacterium), 416, 607 amocontactum (Flavobacterium), 631 Amoebobacter, 16, 23, 26, 848, 849, 850 Amoebomonas, 8, 25, 848 amphibiae (Spirochaeta) , 1065 amphibolus (Vibrio), 702 amplelopsorae (Bacillus), 227 amplus (Micrococcus) , 252 ampullaceus (Micrococcus), 252 amularium (Marmor), 1212 amygdaloides (Bacillus), 648 amylifera (Thiosphaerella) , 998, 999 amyliferum (Spirillum), 217 amyloaceum var. auranticum. (Propioni- bacterium) , 379 amyloaerobius (Bacillus), 738 Amylobacter, 743, 748, 763 amylobacter (Bacillus), 771 amylobacter I (Bacillus), 771, 824 amylobacter II (Bacillus), 771, 824 amylobacter III (Bacillus), 771, 824 amylobacter (Clostridium) , 111 amylobacter (Metallacter), 111 amylobacter immobilis (Bacillus), 790, 826 amylobacter mobilis (Bacillus), 111 amylobacter S and W (Bacillus), 772, 824 amylocella (Vibrio), 203 amyloclasticus (Bacillus) , 813 amyloclasticus intestinalis (Bacillus), 813 amylolactis (Streptococcus) , 325 amylolyticus (Bacillus), 738 amylophilum (Pectinobacter) , 823 amyloruber (Bacillus) , 484 aniyloruber (Erythrobacillus) , 484 amylorubra (Serratia), 484 amylovora (Erwinia), 465, 1135 amylovorum (Achromobacter) , 423 amylovorum (Bacterium), 465 amylovorum (Urobacterium) , 423 amylovorus (Bacillus), 465 amylovorus (Micrococcus), 465 amylozyma (Bacillus) , 771 amylozyme (Clostridium) , 111 amylozxjmicus (Bacillus) , 111 Anaerobacillus , 27, 763 Anaerobe No. Ill, Flligge, 813 Anaerobe No. IV, Flligge, 814 anaerobia (Gaffkya), 284 anaerobic No. V (Bacillus), 818 anaerobic No. VIII (Bacillus) , 813 anaerobicum (Bacterium) , 687, 819 anaerobicus (Bacillus), 820 anaerobicus alcaligenes (Bacillus), 801 anaerobicus caproicus (Bacillus), 820 anaerobicus -liquefaciens (Bacillus), 819 anaerobicus magnus (Streptobacillus) , 823 anaerobicus minutus (Bacillus) , 368 anaerobicus parvus (Coccobacillus), 368 anaerobicus rectus (Streptobacillus) , 822 anaerobicus tenuis (Bacillus), 821 anaerobies (Actinomyces), 922 anaerobies (Oospora), 922 anaerobiontica (Pasteurella) , 581 anaerobium (Achromobacter), 423 anaerobium (Bacterium), 673 anaerobium (Corynebacterium) , 388, 402 anaerobius (Micrococcus), 247, 281 anaerobius (Staphylococcus), 247, 248 anaerobius (Streptococcus), 328, 345 anaerobius (Tetracoccus) , 284 anaerobius carduus (Streptococcus), 328 anaerobius chromogenes (Bacillus), 805 anaerobius diphtheroides (Bacillus) , 402 anaerobius foetidus (Bacillus), 782 anaerobius gonoides (Streptococcus) , 328 anaerobius gracilis (Bacillus), 580 anaerobius haemolysans (Bacillus), 820 anaerobius liquefaciens (Bacillus), 681, 819 anaerobius magnus (Bacillus) , 823 anaerobius major (Staphylococcus) , 281 anaerobius micros (Streptococcus), 328, 330 anaerobius minor (Staphylococcus) , 281 anaerobius perfoetens (Coccobacillus) , 576 anaerobius rectus (Bacillus), 822 anaerobius tenuis (Bacillus) , 821 1396 INDEX OF NAMES OF GENERA AND SPECIES anae-anth anaerobius tenuis (Leptothrix), 365 anaerobius vulgaris (Streptococcus) , 328 anaerobius typ. vulgaris {Streptococcus), 328 anaerogenes (Bacillus), 533 anaerogenes (Bacterium), 533 anaerogenes (Escherichia), 533 Anaeromyces, 925 ananas (Bacillus), 473 ananas (Bacterium), 127, 473 aiianas (Erwinia) 127, 473 ananas (Phytomonas) , 127 ananas (Pseudomonas), 127, 473 Anaplasma, 1100 anata (Escherichia), 523 anatipestifer (Hemophilus), 554 anatipestifer (Pfeifferella) , 554 anatis (Bacterium), 523, 552 anatis (Corynethrix) , 406 anatis (Salmonella), 498,502, 523, 642 (inatis (Spirochaeta) , 1059 unatum (Bacterium), 523 unatum (Salmonella), 43, 523 unatum var. certrycke (Salmonella) , 502 anatum var. muenster (Salmonella) , 523 anatum var. nyborg (Salmonella) , 523 anceps (Bacillus), 648 anceps (Leptotrichia) , 367 anceps (Rasmussenia) , 367 andoi (Vibrio), 201 andropogoni (Bacterium), 142 andropogoni (Phytomonas), 142 andropogoni (Pseudomonas), 142, 169, 1136 anemoeon (Microsporon) , 918 anemones (Galla), 1158 anginosus (Streptococcus), 333, 334 Angiococcus, 1047 angliae (Marmor), 1200 anglomerans (Bacillus), 173 anguillarum (Bacillus) , 673 anguillarum (Bacterium) , 673 anguillarum (Vibrio), 203 angulans (Bacillus), 758 angulata (Phytomonas), 113 angulata (Pseudomonas), 113, 124 angulatum (Bacterium), 113 angulosum (Clostridium), 801, 827 angulosus (Bacillus) , 801, 827 angulosus (Bacteroides) , 801 angustum (Bacterium), 673 anhaemolyticus (Streptococcus), 337 anhaemolyticus vulgaris (Streptococcus), 337, 343 anindolica (Escherichia) , 452 anindolicum (Bacillus), 452 anindolicum (Citrobacter) , 448 annamensis (Corynebacterium) , 402 annamensis (Salmonella), 530 annulare (Photobacterium) , 636 annularis (Microspira) , 636 annulata (Pseudomonas) , 173 annulata (Thiothrix), 990 annulates (Bacterium) , 693 annulatum (Flavobacterium) , 173 annulatus (Actinomyces) , 968 annulatus (Actinomyces) (Streptothrix), 968 amiulatus (Bacillus), 173, 648, 693 annulatus (Micrococcus), 252 annuliformans (Bacillus) , 580 annuliformis (Bacillus) , 738 Annulus, 1212 anodontae (Cristispira), 1055, 1056 onodontae (Spirochaeta), 1055 anolium (Serratia), 461 anoxydana (Colloides), 595 anserina (Borrelia), 1058 anserina (Spirochaeta), 1058 anserina (Spironema), 1058 anserina (Spiroschaudinnia) , 1058 anserina (Treponema), 1058 anserinum (Spirillum), 1058 anserum (Spirillum) , 1058 antarctica (Nitrosospira), 72 antenniforme (Flavobacterium) , 440 antenniformis (Bacillus), 440 antennijormis (Bacterium), 440 anthraciformis (Bacillus) , 648 Anthracillus, 25, 27 anthracis (Aplanobacter), 719 anthracis (Bacillus), 719, 1138 anthracis (Bacillus) (Bacteridiuin), 719 anthracis (Bacillus) (Streptobacter) , 719 1397 anth-arab INDEX OF NAMES OF GENERA AND SPECIES anthracis {Bacterium), 719 anthracis {Pollendera) , 719 anthracis similis (Bacillus) , 738 anthracis symptomatici (Bacillus), 776 anthracis-sym-ptomalici (Clostridium), 11& anthracoides (Bacillus), 648, 716 anthracoides (Bacterium) , 716 Anthrax, 20, 22 anthropopitheci (Spirochaeta) , 1079 antibioticus (Actinomyces), 942 antibioticus (Streptomyces), 942 antirrhini (Bacterium) , 167 antirrhini (Phytomonas) , 167 antirrhini (Pseudomonas) , 167 antirrhini (Xanthomonas), 167 anularis (Cytophaga), 1016 anularius (Bacillus), 648 Apelmocoena, 14, 1032 apertus (Annulus), 1214 aphrophilus (Hemophilus) , 589 aphthicola (Streptococcus) , 337 aphthosum (Bacterium) , 673 aphthosus (Bacillus), 673 apiculatus (Chondromyces) , 1038 apicum (Bacillus) , 648, 738 a2J^^' (Bacterium) , 639 a^Jzi (Phytomonas), 122 apii (Pseudomonas), 122 apiovorus (Bacillus), 470 apis A'^o. 1, A''o. 2 and No. 3 (Bacterium) , 673 apzs (Streptococcus) , 326, 724 apisepticus (Bacillus), 648 Aplanobacter , 8, 705 aporrhoeus (Bacillus), 738 appendicis (Actinomyces) , 922 appendicis (Discomyces) , 922 appendicis (Nocardia) , 922 appendicitis (Bifidibacterium) , 369 aptata (Phytomonas), 114 aptata (Pseudomonas), 43, 114 aptatum (Bacterium), 114 aquamarinus (Achromobacter), 419 aquatile (Flavobacterium), 428, 429 aquatile aurantiacum (Bacterium) , 673 aquatile citreum (Bacterium), 673 aquatile debile (Bacterium) , 673 aquatile flavum (Bacterium), 673 aquatile gasoformans non liquefaciens (Bacterium), 657 aquatile luteum (Bacterium) , 673 aquatile odorans (Bacterium) , 491 aquatilis (Bacillus), 428, 648, 649 aquatilis (Bacterium) , 428, 613 aquatilis (Diplococcus), 694 aquatilis (Micrococcus) , 252, 695 aquatilis (Microspira) , 199 aquatilis (Pseudomonas), 146 aquatilis (Streptococcus), 702 aquatilis (Streptothrix), 976 aquatilis (Vibrio), 199, 632 aquatilis (Zuberella) , 577 aquatilis a (Bacillus), 684 aquatilis albissirnus (Micrococcus) , 695 aquatilis albus (Micrococcus) , 252, 695 aquatilis corninimis (Bacillus), 649, 661, 699 aquatilis communis (Bacterium), 649 aquatilis flavus (Micrococcus), 252 aquatilis gasoformans non liquefaciens (Bacillus), 657 aquatilis invisibilis (Micrococcus), 252 aquatilis magnus (Micrococcus), 695 aquatilis radiatus (Bacillus), 613 aquatilis radiatus (Bacterium) , 613 aquatilis solidus (Bacillus), 658 aquatilis solidus (Bacterium), 658 aquatilis sulcatus (Bacillus), 655 aquatilis sulcatus I (Bacillus), 670 aquatilis sulcatus II (Bacillus) , 670 aquatilis sulcatus III (Bacillus), 669 aquatilis sulcatus IV (Bacillus), 648 aquatilis sulcatus V (Bacillus) , 655 aquatilis -sulcatus -quartus (Bacillus), 649 aquatilis sulcatus quartus (Bacterium) ,649 aquatilis villosus (Bacillus), 671 aqueductum (Leptospira), 1078 aqueum (Bacterium), 759 aqueus (Micrococcus) , 252 aquivivus (Micrococcus) , 695 aquosus (Proactinomyces) , 923 arabinosaceus (Betacoccus), 346 arabinosaceus (Lcuconostoc) , 346 arabinosum (Propionibacterium), 378 1398 INDEX OF NAMES OF GENERA AND SPECIES arab-asce arabinosus {Lactobacillus), 357 arabinotarda Types A and B (Shigella), 544 arachidis (Marmor), 1187 arachnoidea (Beggiatoa), 992, 993 arachnoidea (Oscillaria) , 992 arachnoideus (Bacillus) , 738 araliavora (Erwinia) , 477 araliavorus (Bacillus) , 477 arborescens (Actinomyces), 919 arborescens (Bacillus), 435, 436, 649, 919 arboresceiis (Bacterium), 435 arborescens (Erythrobacillus) , 435 arborescens (Flavobacterium), 429, 435, 439 arborescens (Nocardia) , 919 arborescens lactis (Micrococcus) , 252 arborescens non-liquefaciens (Bacillus), 435 arborescens 7ion-liquefaciens (Bacterium) , 436, 673 arboreus (Bacillus) , 649 Archangium, 1017 archeri (Gafkya), 284 archibaldii (Salmoyiella) , 531 arcticum (Achromobacter) , 423 arcticum (Bacterium) , 673 arechavaleta (Salmonella), 506 arenarius (Bacillus), 738 argenteo-phosphorescens (Bacillus), 634 argenteo-phosphorescens I (Bacillus), 634 argenteo-phosphorescens II (Bacillus) , 634 argenteo-phosphorescens III (Bacillus) , 634 argenteo-phosphorescens (Bacterium) , 634 argenteo-phosphorescens liquefaciens (Ba- cillus), 634 argenteus (Micrococcus) , 252 argentinensis (Spirochaeta) , 1065 argentinensis (Treponema), 1065 argentophosphorescens (Achromobacter) , 634 arguata (Cellulomonas) , 176 arguta (Pseudomonas), 176 aridus (Bacillus), 738 arizona (Salmonella) , 462 Arloingillus, 11, 763 arlongii (Bacillus), 738 armoraciae (Bacillus), 710 aroideae (Bacillus), 474 aroideae (Bacterium) , 474 aroideae (Erwinia), 470, 474, 1129, 1136 aroideae (Pectobacterium) , 474 aromafaciens (Achromobacter) , 423 aromafaciens (Bacterium) , 423 aromatica (Pseudomonas) , 146 aromatica var. quercitopyrogallica (Pseu- domonas), 146 aromaticum (Flavobacterium) , 457 aromaticus (Bacillus), 434, 457, 649, 743 aromaticus (Bacterium), 457 aromaticus (Streptococcus) , 337 aromaticus butyri (Bacillus), 440 aromaticus lactis (Bacillus) , 434 arthritica (Micrococcus) , 301 arthritica (Neisseria), 301 arthritidis (Bacterium), 674 nrthritidis (Murimyces) , 1292 arthritidis chronicae (Bacillus) , 674 arthritidis-muris (Corynebacteriurn), 402 Arthrobacter, 7 Arthrobactridium , 7 Arthrobactrillium , 7, 82 Arthrobactrinium , 7 , 82 Arthromitus, 1003 Arthrostreptokokkus, 312 arthrotropicus (Musculomyces) , 1243 arthuri (Bacillus) , 639 articulata (Pseudomonas) , 146 arliculatum (Bacterium) , 759 articular mn (Streptococcus) , 337 artus (Phagus), 1133 arvalis (Grahamella) , 1109 arvicolae (Bartonella) , 1104 arvicolae (Haemobartonella) , 1104 arvilla (Pseudomonas), 105 arvillum (Achromobacter), 105 asaccharolyticus (Micrococcus), 246 asaccharolyticus (Staphylococcus) , 246 asaccharolyticus var. indolicus (Staphylo- coccus), 247, 264 asalignus (Streptococcus), 337 ascendens (Acetobacter), 43, 185, 692 ascendens (Bacterium), 185 ascendens (Ulvina), 692 1399 asci-aura INDEX OF NAMES OF GENERA AND SPECIES ascitis (Corytiebacteriuni), 402 Ascococcus, 6, 235 ascojormans {Botryococcus), 253 ascoformans (Micrococcus) , 252, 253 ascoforrnans (Staphylococcus), 253 ascoformis (Micrococcus) , 253 asiaticum (Bacterium) , 450 asiaticus (Bacillus), 450, 73S asiaticus (Proteus), 450 asiaticus (Salmonella) , 450 asiaticus mobilis (Bacillus) , 450 asiaticus mobilis (Salmonella), 450 asparagi (Bacillus) , 759 asparagi (Bacteriuyn) , 759 asper (Micrococcus) , 253 asplenii (Phytomonas) , 696 asporiferum. (Bacterium) , 674 assimilis (Bacillus) , 649 assurgens (Archangium) , 1019 Asiasm, 20, 22, 705 asteracearum. (Bacillus) , 477 asteracearum (Erwinia), 477 asteriformis (Bacillus), 612 asteris (Bacillus) , 738 Asterococcus, 1289, 1291 asteroide (Leptothrix) , 218, 365 asteroide (Mycobacterium) , 896 .Is^eroides, 892 asteroides (Actinomyces), 896 aster oides (Asteroides) , 896 asteroides (Cladothrix) , 896 asteroides (Discomyces) , 896 asteroides (Nocardia), 896, 897, 918 asteroides (Oospora), 896 asteroides (Proactinomyces ) , 896 asteroides (Strepiotrix) , 896 asteroides var. crateriformis (Nocardia), 897 asteroides var. crnteriformis (Proactino- myces), 897 asteroides var. decolor (Nocardia), 897 asteroides var. decolor (Proactinomyces), 897 asteroides var. gypsoides (Nocardia) , 897 asteroides var. gypsoides (Proactino- myces), 897 asteroides var. serratus (Actinomyces), ■ 917 Asteroniyces, 1291 asterospora (Astasia), 720 asterosporus (Aerobacillus), 720 asterosporus (Bacillus) , 720, 748 asterosporus alpha (Bacillus) , 720 astheniae (Bacillus), 448 astheniae (Bacterium) , 448 astheniae (Escherichia) , 448 asthenoalgiae (Leptospira), 1078 asthenogenes (Bacillus) , 738 astragali (Bacterium), 139 astragali (Phytomonas) , 139 astragali (Pseudomonas), 139 astri (Marmor), 1196 astrictum (Marmor), 1165. 1167 astrictum var. aucuba (Marmor) , 1168 astrictum var. chlorogenus (Marmor), 1168 astrictus (Phagus), 1133 aterrimus (Bacillus), 711 aterrimus tschitensis (Bacillus) , 738 atherton (Salmonella), 701 atlantica (Pseudomonas), 697 yliremis, 13 atrofaciens (Bacterium), 120 atrofaciens (Phytomonas), 121 atrofaciens (Pseudomonas), 120 atroseptica (Erwinia), 468, 470, 1131 atrosepticus (Bacillus), 468, 470 attenuatum (Spirillum) , 43, 217 attenuatum (Spirosoma) , 217 atypica pse udot uherk ulosa (A ct inom yces ) , 973 aucuba (Marmor), 1175 aucuba var. canadense (Marmor), 1175 auciibicola (Pseudomonas), 146 aurantia (Spirochaeta) , 1053 aurantiaca (Cladothrix), 896 aurantiaca (Cytophaga), 1013 aurantiaca (Merismopedia) , 251, 290 aurantiaca (Nocardia), 896 aurantiaca (Oospora), 896 aurantiaca (Paulosarcina) , 288 aurantiaca (Sarcina), 243, 288 aurantiaca (Stigmatella), 1037 aurantiaca (Streptothrix) , 896 1400 INDEX OF NAMES OF GENEIL\ AND SPECIES aura-avis aurantiac ilm (Agarbacterium), 630 aurantiacum (Bacteridium) , 243 aurantiacurn (Chromobacterium) , 440 aurantiacum (CorT/nebacterium) , 402, 404 aurantiacum (Flavobacterium) , 440 aurantiacum (Polycephalum) , 1037 aurantiacus (Actinomyces), 896 aurantiacus (Aurococcus), 243 aurantiacus (Bacillus), 440 aurantiacus (Bacleriujn) , 440 aurantiacus (Chondromyces), 1037, 1038 aurantiacus (Micrococcus), 243, 265, 275, 290 aurantiacus (Pediococcus) , 243, 290 aurantiacus (Staphylococcus) , 243 aurantiacus (Streptococcus) , 243, 253, 337 aurantiacus-sorghi (Micrococcus), 243, 253 aurantiacus tingitanus (Bacillus), 145, 646 aurantiacus var. frutescens (Chondro- myces), 1037, 1039 aurantibutyricum (Clostridium), 819 aurantii (Bacillus), 674 aurantii (Bacterium) , 674 aurantinum (Flavobacterium), 440 aurantinus (Bacillus) , 4i0 aurantium (Plocamobacterium) , 674 aurantium roseum (Bacterium) , 392, 674 aurantius (Bacillus), 649, 738, 739 aurantius (Cellulomonas), 739 aurea (Actinomyces), 943, 968 aurea (Nocardia), 968 aurea (Oospora) , 968 aurea (Pseudomonas) , 146, 697 aurea (Sarcina), 290 aurea (Streptothrix), 968 aureo-flavus (Bacillus), 674 aureo flavus (Bacterium), 674 Aureogenus, 1154 aurescens (Bacillus), 440, 474, 674 aurescens (Bacterium), 440, 445, 484, 674 aurescens (Flavobacterium) , 440 aurescens (Sarcina), 290 aurescens var. mucosa (Sarcina), 290 aureum (Bacterium) , 674 aureum (Polyangium), 1031 aureum (Spirillum), 203 aureum (Spirosoma), 203 aureus (Actinomyces), 943, 968, 977 aureus. (Aurococcus) , 241 aureus (Bacillus), 649, 674, 716 aureus (Flexibacter) , 38 aureus (Micrococcus) , 241, 251, 252, 253, 254, 256, 258, 265, 266, 268, 269, 271, 276, 279 aureus (Myxobacter), 1026 aureus (Staphylococcus), 241, 1140, 1141 aureus (Streptomycesj, 943 aa/-ews (F/6no), 203, 204 aureus var. eguz (Staphylococcus) , 253 aureus lactis (Micrococcus) , 253 aureus minutissiinus (Bacillus) , 663 aureus sarciniformis (Staphylococcus), 281 oi/r«s (Bacillus), 402 awr/s (Corynebacterium), 402 .4.«rococcus, 8, 235 aurogenes (Bacillus), 617 aurogenes (Cellulomonas), 617, 622 aurogenes var. ai6ws (Bacillus) , 622 australiense (Lethum), 1223 australiense var. lethale (Lethum), 1224 australiense var. typicvm (Lethum), 1224 australiensis CChlorogenus), 1147 autumnalis (Leptospira), 1078 autumnalis (Spirochaeta), 1078 autumnalis A (Spirochaeta), 1078 autumnalis Type B (Leptospira), 1077, 1078 auxinophilum (Bacterium), 145 auadi (Actinomyces), 915, 924 avenae (Bacillus), 116 avenae (Fractilinea), 1162 avenae (Phytomonas) , 116 avenae (Pseudomonas), 116 avicida (Pasteurella), 547, 552, 642 avicidum (Bacterium), 546, 547 avicidus ' Coccobacillus) , 547 avidum (Corynebacterium) , 388, 402 avidus (Bacteroides) , 380, 402 avidus (Vibrio), 702 aviseptica (Pasteurella), 523, 547 avisepticum (Bacterium), 547 avisepticus (Bacillus), 530, 547 1401 aviu-bact INDEX OF NAMES OF GENERA AND SPECIES avium (Bacillus), 400 avium (Bacterium) , 674 avium (Borreliota), 1229 avium (Mycobacterium), 878, 879, 880 avium (Pasleurdla) , 547 avium (Rickelisia), 1095 avium (Strongyloplasma), 1229 avium (Tarpeia), 1274 azolicus (Bacillus), 772, 824 Azotobacter, 19, 21, 26, 29, 31, 216, 219, 221 azotobacter (Bacillus), 219 azotogcna (Pseudomotias) , 697 Azotomonas, 8, 219, 221 azureus (Bacillus), 049 babcsi (Bacillus), 649 babesi (Neisseria), 254, 696 Babesia, 312 babesii (Bacterium), 674 baccarinii (Bacillus), 466 baccarinii (Clostridium), 478 baccatus (Micrococcus) (Sarcina.) , 253 bacillifera (Pelogloea) , 871 bacilliformis (Bartonella), 1101, 1105 bacilliformis (Bar Ionia) , 1101 bacillosum (Amoebobacter), 849 Bacillus, 6, 7, 13, 15, 18, 19, 22, 27, 30, 31, 42, 43, 46, 63, 76, 179, 632, 643, 704, 705, 763, 1008 Bacillus I, Bienstock, 755 Bacillus II, Bienstock, 753 Bacillus II, Leube, 668 Bacillus a, Guillebeau, 662 Bacillus a, b, c, d, c, /, h and i, Vignal, 647 Bacillus A, Grigoroff, 354 Bacillus A, Maggiora, 671 Bacillus B, Hoffmann, 749 Bacillus B, Maggiora, 661 Bacillus D, Foutin, 744 Bacillus D, Peters, 741 Bacillus G, Maggiora, 650 Bacillus H, Maggiora, 662 Bacillus X, Moore and White, 726 Bacillus a, Busse, 639 Bacillus a, von Freudenreicb, 356 Bacillus y, von Freudenreicb, 358 Bacillus 8, von Freudenreicb, 360 Bacillus e, von Freudenreicb, 352 Bacillus No. 18, Conn, 760 Bacillus No. 25, Conn, 753 Bacillus No. 1^1, Conn, 423 Bacillus No. 2, Fulles, 668 Bacillus No. 2, Kedrowski, 814 Bacillus No. 3, Pansini, 718 Bacillus No. 6, Pansini, 710 Bacillus No. 8, Pansini, 738 Bacillus No. XVI, Adametz, 718 Bacilhis No. XVII, Adametz, 760 Bacillus No. I, Fliigge, 716, 725 Bacillus No. II, Fliigge, 741 Bacillus No. Ill, Fliigge, 738 Bacillus No. IV, Fliigge, 744 Bacillus No. V, Fliigge, 716 Bacillus No. VI, Fliigge, 743 Bacillus No. VII, Flugge, 747 Bacillus No. VIII, Fliigge, 748 Bacillus No. IX, Fliigge, 710 Bacillus No. X, Fliigge, 709 Bacillus No. XI, Flugge, 743 Bacillus No. XII, Fliigge, 755, 761 Bacillus of swine plague, 508 Bacillus sp., Sordelli, 777 Bacteridium, 6, 43, 705 Bacterienart No. 12, Lembke, 745 bacterifera (Cylindrogloea), 874 Bacteriophagum, 1128, 1129 bacteriophagus (Protobios), 1129 Bacteriopsis, 179, 365 Bacterium, 5, 7, 10, 13, 15, 17, 18, 19, 28, 30, 37, 42, 46, 76, 82, 179, 435, 596, 597, 599, 612, 694, 705, 763 Bacterium A, Peters, 657 Bacterium B, Peters, 664 Bacterium of hog cholera, 508 Bacterium of swine plague, 508 Bacteroidea, 20 Bacteroides, 22, 23, 27, 31, 32,218,365, 564, 763, 1296 Bactoderma, 76 Bactrella, 705 Bactridiurn, 7, 705, 763 Bactrillius, 83 Bactrillum, 7, 13, 25, 27, 82, 705 1402 INDEX OF NAMES OF GENERA AND SPECIES bact-bell Bactrinium, 7, 82, 705 Bactrinius, 83 badius (Bacillus), 739 badius (Micrococcus), 253 bahiensis (Actinomyces) , 919 bahiensis (Discomyces) , 919 bahiensis (Nocardia), 919 bahiensis (Oospora), 919 balaenae (Clostridium) , 775, 819 balanitidis (Spirochaeta) , 1065 balanitidis (Spironema), 1065 balanitidis (Spiroschaudinnia) , 1065 balanitidis (Treponema) , 1065 balbianii (Bacillus) , 674 balbianii (Bacterium), 674 balbianii (Cristispira), 1055, 1056, 1057 balbianii (Spirochaeta) , 1055 balbianii (Trypanosoma), 1055 balcanus (Bacillus), 739 balfourii (Grahamella) , 1109 Balkanella, 10 ballerup (Salmonella) , 529 balticum (Phofobacterium) , 635, 636 balticus (librio), 636 balustinum (Flavobacterium), 437 bamptonii (Chromobacterium) , 234 bantam (Salmojiella) , 523 barati (Inflabilis) , 823 barbareae (Phytomonas) , 154 barbareae (Xanthomonas), 154 barbatum (Bacterium) , 761 barbitistes (Bacillus), 739 baregensis purpureus (Micrococcus), 253 bareilly (Salmonella) , 511 bareilly var. mikawasima (Salmonella). 511 ' barentsianum (Bacterium), 674 barker i (Bacillus), 129 barkeri (Bacterium) , 129 barkeri (Phytomonas), 129 barkeri (Pseudomonas), 129, 134 Bartonella, 37, 1100, 1102, 1109 Bartonella sp., 1108 Bartonia, 1100 batatae (Bacilhis), 739 batavia (Salmonella) , 524 bataviae (Leptospira), 1078 batrachorum (Arthromitus), 1003 batrachorum (Bartonella) , 1108 batrachorum (Haemobartonella) , 1108 balrochorum (Micrococcus), 1123 bauri (Bacterium), 108 beaufortensis (Pseudomonas) , 697 beckii (Bacterium) , 552 beddardii (Actinomyces), 963 beddardii (Streptomyces) , 963 Beggiatoa, 12, 16, 18, 19, 24, 26, 42, 988, 990, 993, 994, 1007 beggiatoides (Oscillatoria) , 992 begoniae (Bacterium) , 155 begoniae (Phytomonas), 155 begoniae (Xanthomonas), 155 beigeliana (Zoogloea), 253 beigelianum (Sclerotium) , 253 beigelli (Chlamydatomus), 253 beigelii (Hyalococcus) , 253 beigelii (Micrococcus) , 253 beigelii (Pleurococcus) , 253 beigelii (Trichosporum) , 253 beijerincki (Bacillus) , 356, 650, 691 beijerincki (Bacterium) , 674 beijerinckii (Azotobacter) , 219 beijerinckii (Clostridium), 772 beijerinckii (Lactobacillus), 357 beijerinckii (Pseudomonas), 109 beijerinckii (Rhizobium), 225 beijerinckii (Rhizomonas) , 225 beijerinckii (Sarcina), 286 beijerinckii (Thiobacterium) , 81 beijerinckii (Urobacillus) , 691 beijerinckii (Vibrio), 203 beijerinckii var. jacobsenii (Thiobac- teriinn), 81 bekkerii (Mycobacterium) , 890 6eZ/ar?/^ (Bacillus), 649, 803 belfantii (Clostridium), 803 belfantii (Endosporus) , 649, 803 belfastiensis II (Bacillus), 533 belfastiensis V (Bacillus) , 534 belfastiensis (Bacterium) , 533 belfastiensis (Eberthella) , 533 bcllisari (Actinomyces), 968 bellonensis (Bacilhis), 777, 825 bellonensis (Clostridium) , 778 1403 bell-bife INDEX OF NAMES OF GENERA AND SPECIES hellus {Bacillus), 739 belorinensis (Bacillus), 360 bemisiae (Ruga), 1219 benlotensis (Bacillus), 533 bentotensis (Bacterium) , 533 benlotensis (Castellanus) , 533 bentotensis (Eberthella) , 533 benzoli (Bacillus), 649 benzoli a and b (Bacterium) , 674 berardinisi (Discornyces) , 918 berardinisi (Nocardia), 918 berbera (Borrelia), 1061 berhera (Spirochaeta) , 1061 berbera (Spirone?na), 1061 berbera (Spiroschaudinnia) , 1061 berberidis (Bacterium), 115 berberidis (Phytomonas), 115 berberidis (Pseudomonas), 115 berberum (Treponema), 1061 berestneffi (Nocardia), 919 berestneffii (Actinomyces, 919 berestneffii (Discornyces), 919 bereslnewi (Bacillus), 918 bergerac (Annulus), 1213, 1216 beri-beri (Micrococcus), 253 beribericus (Bacillus), 649 bernardinisi (Actinomyces), 918 bernensis (Bacillus) , 739 berolinensis (Bacillus), 234, 360, 650 berolinensis (Lactobacillus) , 360 berolinensis (Microspira) , 196 berolinensis (Mycobacterium) , 888, 890 berolinensis (Pseudomonas) , 697, 698 berolinensis (Vibrio), 196, 204 berolinensis fasciformis (Saccharo- bacillus), 360 berolinensis indicus (Bacillus) , 697 berolinensis indicus (Bacterium), 697 6erto (Salmonella), 518 bertherandi (Coniothecium) , 289 besseri (Bacterium) , 674 bessoni (Kurthia) , 613 be^a (Bacillus), 650, 739 6e/a (Micrococcus), 269 6e^a (Nocardia), 976 beta (Phagus), 1141 beta (Scelus), 1237 6e/a (Streptothrix) , 976 beta (Tarpeia), 1270 Betabacterium, 9, 30, 350 Betacoccus 9, 30, 346 betadelbrueckii (Lactobacillus), 363 6e/ae (Bacillus), 477, 639 6e/ae (Bacterium), 144, 639 fee/ae (Butijlobacter) , 781, 825 betae (Corium), 1204 betae (Marmor), 1178 6eep//c (Bacterium), 130 /,e/Zis (Phytomonas), 130 betlis (Pseudomonas), 130 biacutum (Bacillus), 739 biacutum (Fusobacterium), 582 biacutus (Fusiformis), 582 biazotea (Cellulomonas), 617 biazoteus (Bacillus), 617 bibula (Cellulomonas), 615 bibulum (Bacterium), 615 bibulus (Bacillus), 615 bicolor (Actinomyces), 919 bicolor (Micrococcus), 254 bicolor (Nocardia), 919 bicolor (Sarcina), 290 bienstocki (Putrificus), 799 bienstockii (Bacillus), 544 bienstockii (Bacterium), 544 bienstockii (Eberthella) , 544 bienstockii (Shigella), 544 bifermentans (Bacillus) , 787 1404 INDEX OF NAMES OF GENERA AND SPECIES bife-bore bifermentaus (Clostridium), 309, 782, 787, 818, 825 bifermentans {Martellillus) , 787 hifermentans sporogenes (Bacillus), 787 bifida (Lieskeella), 986 bifida {Nocardia), 353 Bifidibacterium, 34, 38, 369 Bifidobacterium, 349, 369 bifidum {Bacterium) , 354 bifidum {Bifidibacterium) , 353, 369 bifidus {Actinomyces), 353 bifidus {Bacillus), 353 bifidus {Bacteroides) , 353, 354 bifidus {Cohnistreptothrix), 353 bifidus (Lactobacillus), 353, 354. 369 bifidus II {Lactobacillus), 354 bifidus aerobius {Bacillus) , 361 bifidus capitatus {Bacillus) , 361 bifidus communis {Bacillus) , 353, 36! bifilaris {Lieskeella) , 986 biflexa (Leptospira), 1077, 1078 biflexa {Spirochaeta), 1077 bijorme {Eubacterium), 368 biformis {Bacteroides), 362, 36S bifurcatum {Bifidibacterium), 369 bifurcatus gazogenes {Bacillus), 369 biliohemoglohinuriae {Leptospira), 107S bilio-hemoglohinuriae {Spirochaeta), 1078 Billetia, 6S0 billingsi {Bacillus) , 650 billrothii {Ascococcus) , 254 billrothii {Micrococcus), 254 binucleatum {Bacterium), 760 bipolare multocidum {Bacterium), 547 bipolaris {Bacillus), 548, 718 hipolaris br)visepticus {Bacillus), 547 bipolaris bubalisepticus {Bacillus), 548 bipolaris caprisepticus {Bacillus) , 553 bipolaris ovisepticus {Bacillus), 554 bipolaris plurisepticus {Bacillus) , 546 bipolaris septicus {Bacillus) , 546, 547 bipunctata (Macromonas) , 997, 1001 bipunctata {Pseudomonas) , 997, 1001 bipunctata (Thiospira), 212 bipunctatum {Spirillum) , 212 biskra {Micrococcus), 254 biskrae {Staphylococcus), 254 bispebjerg {Salmonella), 506 bizzozerianus {Bacillus), 743 blackwellii {Actinomyces), 910 blackwellii (Nocardia), 910 blanci {Rickettsia), 1088 blarinae {Grahamella) , 1110 blarinae (Haemobartonellai, 1107 blasticus (Choiidrococcus). 1008, 1046 Blastocaulis, 35, 836 blattellae (Corynebacterium), 402 blegdam {Salmonella) , 518 bleischii {Bacillus), 533 boas-oppleri {Lactobacillus), 352 bobiliae {Actinomyces) , 937 bobiliae (Streptomyces), 937 6oZe/i {Micrococcus), 254 boletus (Melittangium), 1006, 1034 bollingeri (Pasteurella), 547 bolognesii-chiurcoi {Actinomyces) , 915 bolognesii-chiurcoi (Malbrachea), 915 bombycis {Aerobacter) , 490 bombycis {Bacillus) , 650, 739 bombycis {Bacterium,) , 650 bombycis (Borrelina), 1226 bombycis {Chalmydozcon), 1226 bombycis {Diplococcus), 336 bombycis {Micrococcus), 254 bombycis (Microzyma), 254 bombycis {Nosema), 254 botnbycis {Proteus), 490 bombycis {Streptococcus), 254, 265, 337 bombycis non-liquefaciens {Bacillus) , 739 bomhycivorum {Bacterium), 490 bombycoides {Bacillus), 739 l)omby septicus {Bacillus), 739 bonariensis (Leptospira) , 1078 bonariensis (Salmonella), 514 bonhoffii (Microspira), 202 bonvicini {Streptococcus) , 337 bookeri (Alcaligenes), 415, 416 bookeri (Bacillus), 415, 650 bookeri (Bacterium), 415 borbeck (Salmonella) , 527 borborokoites {Bacillus), 739 bordonii (Bacterium) , 691 bordonii {Klebsiella), 691 boreale (Bacterium), 625 1405 bore-bras INDEX OF NAMES OF GENERA AND SPECIES boreale (Flavobacterium) , 625 boreopolis (Pseudomonas), 94 boreus (Micrococcus), 254 bornensis (Erro), 1256 Borrelia, 19, 34, 35, 37, 42, 1057, 1058 Borrelina, 1225 Borreliota, 1229 Borrelomyces, 1291 borstelensis (Bacillus), 739 bosporuni (Bacterium), 145 bossonis (Bacterium) , 675 bostroemi (Actinomyces), 970 botkini (Bacillus), 813 botryogenus (Micrococcus) , 253 Botryomyces, 235 Botulinea, 20 botulinum (Clostridium), 778, 784 botulinum D (Bacillus) (Clostridium) , lid botulinum Type A (Clostridium) , 784 botulinum Type B (Clostridium) , 784 botulinum Type C (Clostridium), 779 botulinum Type D (Clostridium) , 779 botulinum Tj'pe E (Clostridium), 779 Botulinus, 20, 22, 763 botulinus (Bacillus) , 778 botulinus (Ermengemillus) , 778 botulinus Type C (Bacillus) , 779 Botulobacillus , 8, 763 bouffardi (Pasteurella) , 553 boutrouxii (Bacillus) , 675 boutrouxii (Bacterium), 675 bovicida (Bacterium), 548 bovidae (Treponema) , 1076 Bovimyces, 1291 bovina (Listerella) , 409 bovinum (Scelus), 1239 bovinus (Micrococcus) , 254, 337 bovinus (Streptococcus), 337 bovis (Actinomyces), 925, 926, 927 bows (Bacillus), 652 ftowi's (Bacterium) 675 boOTS (Bartonella), 1106 6ovjs (Betacoccus) , 347 6oOTS (Cladoihrix) , 925 bovis (Coryuebacterium), 390, 391 6orzs (Corynethrix) , 401 &o?iis (Discomyces) , 925 /boot's (Ehrlichia), 1095 6oOTS (Grahamella) , 1110 6cwis (Haematococcus) , 254 bovis (Haemobartonella), 1106 bow's (Hemophilus), 591 6ot'is (Leptospira), 1078 &OOTS (Leuconostoc) , 347 6ovzs (Micrococcus), 254 bovis (Molitor), 1242 bovis (Moraxella), 591 bofi's (Nocardia), 925 boufs (Oospora), 925 6oris (Proactinomyces) , 925 6ovzs (Rickettsia), 1095 feovi's (Sphaerotilis) , 925 bow's (Staphylococcus), 264, 281 bovis (Streptococcus), 320, 321, 322 boCTS (Streptothrix) , 925 bovis (Tortor), 1276 bom var. nigerianus (Actinomyces), bovis albus (Actinomyces) , 968 bovis-caffris (Spirochaeta) , 1065 bovis-caffris (Spironema), 1065 bori's communis (Streptothrix) , 925 bori's farcinicus (Actinomyces) , 895 bow's luteoroseus (Actinojnyces) , 971 bow's morbificans (Bacillus) , 514 bovis-morbificans (Salmonella) , 514 boi/'i's sulfureus (Actinomyces) , 925 boviseptica (Pasteurella), 547 bovisepticus (Bacillus), 547 bovisepticus (Bacterium), 547 bovium (Pasteurella), 547 bowlesiae (Pseudomonas), 125 bowlesii (Bacterium), 125 bowlesii (Phytomonas) , 125 Brachybacterium, 312, 349 brachysporum (Bacterium), 759 brachythrix (Bacillus), 650 braenderup (Salmonella) , 511 brandenburg (Salmonella) , 505 brandenburgensis (Salmonella) , 505 brandenburgiensis (Bacillus), 726 brandti (Bacterium) , 108 branhamii (Micrococcus) , 303 brasiliensis (Actinomyces), 918 hrasiliensis (Discomyces), 918 1406 INDEX OP NAMES OF GENERA AND SPECIES bras-bucc brasiliensis (Escherichia) , 452 brasiliensis (Nocardia), 918 brasiliensis (Oospora), 918 brasiliensis (Rickettsia), 1087 brasiliensis (Streptothrix) , 918 brassicae (Bacillus), 675, 714, 718 brassicae (Bacterium), 357, 675, 714, 718 brassicae (Borrelina), 1227 brassicae (Lactobacillus), 357 brassicae (Marmor), 1177 brassicae acidae (Bacterium), 146, 675 brassicae acidae (Pseudomonas) , 146 brassicae fermentatae (Bacillus), 358 brassicaevorus (Bacillus) , All bredemannii (Bacillus), 739 bredeney (Salmonella), 507 breitfussi (Bacterium), 675 breslau (Salmonella), 502 breslaviensis (Bacillus), 502 breslaviensis (Bacterium) , 502 breve (Bacterium), 439 breve (Betabacteritim) , 358 breve (Flavobacterium), 439 brevis (Bacillus), 439, 725, 739, 744 brevis (Lactobacillus), 358, 359, 361, 363 brevis (Streptococcus) , 337, 696 brevis o (Bacillus), 716 brevis non hemolyiicus (Streptococcus), 338 hrevis var. rudensis (Lactobacillus) , 357. 359 brevissimum (Bacterium), 675 Brevistreptothrix, 892, 925 Briareus, 1233 briensis (Nitrospira), 72 brightii (Streptococcus) , 338 briosianum (Bacterium) , 639 briosii (Bacterium), 145 bronchiale (Treponema), 1065 bronchialis (Actinomyces), 922 bronchialis (Discomyces), 922 bronchialis (Oospora), 922 bronchialis (Spirochaeta) , 1065 bronchialis (Spiroschaudinnia) , 1065 bronchicanis (Bacillus) , 562 bronchicanis (Bacterium), 562 bronchiseptica (Brucella), 562 bronchisepticus (Alcaligenes) , 416, 562 bronchisepticus (Bacillus), 562 bronchisepticus (Bacterium) , 562 bronchitica (Anaeromyces) , 926 bronchitica (Cohnistrepiothrix) , 926 bronchitidis (Bacillus) , 739 bronchitidis putridae (Bacillus) , 739 bronchopneumoniae (Ehrlichia), 1118 bronchopneumoniae (Miyagawanella) , 1118 Brucella, 17, 26, 32, 42, 43, 560, 562 Brucella melitensis var. melitensis, 561 brumptii (Grahamella) , 1110 bruneum (Corynebacterium), 403 bruneum y arborescens (Bacterium), 403 bruneus (Bacillus), 740 bruni (Actinomyces), 923 bruni (Discomyces) , 923 /jrunz (Nocardia), 923 brunneoflavum (Bacterium) , 675 brunneoflavus (Bacillus), 675 brunneum (Bacteridium), 650 brunneum (Bacterium) , 650, 680, 737, 740 brunneum (Flavobacterium), 440 brunneus (Bacillus), 440, 650, 737, 740 brunneus (Micrococcus), 675 brunneus rigensis (Bacillus), 430 bruntzii (Bacillus), 644, 645 bruntzii (Serratia), 644 bubalorum (Clostridium) , 118, 825 bubalseptica (Bacillus), 548 bubalseptica (Pasteurella) , 548 hucallis (Bacterium) , 440 buccale (Bacterium) , 365, 440 buccale (Borrelia), 1062 buccale (Spirillum), 1062 buccale (Spironema), 1062 buccale (Treponema), 1062 buccalis (Actinomyces) , 922 huccalis (Ascococcus), 693 buccalis (Bacillus), 365, 650 buccalis (Discomyces), 922 buccalis (Flavobacterium), 440, 647 buccalis (Leptospira), 1079 buccalis (Leptothrix) , 365, 366 buccalis (Leptotrichia), 364, 365 buccalis (Micrococcus), 329 1407 bucc-buty index of names of genera and species buccalis (Microspira), 1062 buccalis (Molitor), 1242 buccalis {Nocardia), 922 buccalis (Oospora), 922 buccalis (Rastnussenia) , 365 buccalis (Spii'ochaeta), 1062, 1065 buccalis {Spiroschaudinnia), 1062 buccalis (Streptococcus), 338 buccalis {Streptothrix) , 923 buccalis (Syncrotis) , 365 buccalis (Vibrio), 203 buccalis fortuitus (Bacillus) , 647, 650 buccalis fortuitus (Bacterium) , 650 buccalis minutus (Bacillus), 440 buccalis minutus (Bacterium) , 440 buccalis muciferens (Bacillus) , 650 buccalis septicus (Bacillus) , 650 bucco-pharyngei (Spirochaeta) , 1065 bucco-pharyngei (Treponema), 1065 buchneri (Bacillus) , 359 })uchneri (Lactobacillus), 359, 695 buchneri (Ulvina), 695 budapest (Salmonella), 505 budayi (Bacillus), 791 hufo (Agarbacterium), 628 bufonis (Spirochaeta), 1066 bufonis (Spironema), 1066 bufonis (Spiroschaudinnia), 1066 bufonis (Treponema), 1066 /;uZ6osa (F{6no), 203 buhjaricum (Acidobacterium) , 354 l)u.Ujaricum (Bacterium), 354, 687, 695 bulgaricum (Plocamobacterium) , 354 bulgaricutn (Thermobacterimn) , 354 bulgaricus (Bacillus), 354 bulgaricus (Lactobacillus), 354, 362, 364, 695 bullata (Mj'coplana), 191 hullosum (Bacterium), 675 bullosus (Bacillus), 580 bullosus (Bacteroides) , 580 bullosus (Spherocillus) , 580 burchardti (Micrococcus), 254 burgeri (Bacillus), 116 burneti (Rickettsia), 1092 burneti (Rickettsia) (Coxiella), 1092 burneti var. americana (Rickettsia), 1092 burnetii (Coxiella), 1090, 1092 6w?Ti (Bacillus) , 726 busae asiaticae (Bacterium), 357 bus aeasiaticus (Lactobacillus), 357 bussei (Bacillus), 477 bussei (Erwinia), 477 butantan (Salmonella), 524 butlerovii (Bacillus), 740 butschlii (Bacillus), 740, 742, 744 butylaceticum (Bacillus), 781, 825 Butylbacillus, 771 butylicum (Amylobacter) , 771, 813 butylicum (Clostridium), 771 butylicum (Granulobacter) , 771, 824 butylicus (Bacillus), 680, 771, 824 butylicus B. F. (Bacillus), 781, 825 B utylobacter , 763 butyri (Achromobacter), 421 Butyribacillus, 8, 763 6 «^?/ri (Bacillus) , 650 butyri I (Bacillus), 650 butyri II (Bacillus), 659 Butyribacterium, 368, 380, 402 butyri (Diplococcus) , 254 butyri (Flavobacterium) , 440 butyri (Mycobacterium), 890 butyri (Micrococcus), 254 butyri (Pseudomonas), 146, 697 butyri aromafaciens (Bacillus), 421 butyriaromafaciens (Bacterium), 421 butyri -aromafaciens (Micrococcus), 421 butyri colloideum (Bacterium) , 676 butyri fluorescens (Bacillus) , 146 butyri fluorescens (Bacterium), 697 butyri jluorescens (Micrococcus), 254 butyrica (Botulinea), 22 butyrica (Sarcina), 290 Butyriclostridixim , 11, 763 butyricum (Bactridium) , 111, 819, 824 butyricum (Clostridium), 716, 770, 771. 772, 781, 813, 824, 825 butyricum (Mycobacterium), 888, 890 butyricum I (Clostridium), 771 butyricum II (Clostridium), 771 butyricum III (Clostridium), 111 butyricum iodophilum (Clostridiu7n), 772, 824 1408 INDEX OF NAMES OF GENEEA AND SPECIES buty~cana hutyricum var. americanum {Clostri- dium), 819 butyricus {Bacillus), 716, 727, 740, 770, 813, 820 butyricus {Micrococcus), 254, 338 butyricus {Streptococcus), 338 Irutyricus {Tetracoccus) , 254 butyricus asporogenes ijnmobilis {Bacil- lus), 790 butyricus diynorphus {Bacillus) , 813 butyricus putrefaciens {Bacillus) , 799 Butyrisarcina, 29, 30, 31, 285 byzantinea {Brucella) , 693 byzantineum {Coccobacterium) , 693 cacaoi {Actinomyces), 951 cacaoi (Streptomyces), 951 Cacospira, 12, 13, 28, 1058 cacticida {Erwinia), 478 cacticidus {Bacillus) , 478 cactivorum {Bacterium) , 613 cadaveris {Bacillus), 669, 675, 791, 799 cadaveris {Bacterium,), 675, 791 cadaveris {Clostridium), 791 cadaveris {Eubacterium) , 367, 791 cadaveris {Pleciridium) , 799 cadaveris {Streptococcus) , 338 cadaveris butyricum {Bacterium), 791 cadaveris butyricus {Bacillus), 367, 380, 791, 826 cadaveris grandis {Bacillus), 813 cadaveris sporogenes {Bacillus) , 799 cadaveris sporogenes (anaerobicus) {Ba- cillus), 799, 826 Caduceus, 33, 34, 763 caducus (Phagus), 1141 caeci {Bacillus), 650 caesia (Cellulomonas), 619 caesirae retortiformis {Spirochaeta) , 1066 caesirae septentrionalis {Spirochaeta), 1066 caesius {Bacillus), 619 cajae {Coccobacillus) , 690 cajMS {Bacillus), 690 calceum {Ferribacterium) , 834 calceum {Siderobacter) , 834 calceus {Bacillus), 834 calciphila {Pseudomonas) , 146 calciprecipitans (Pseudomonas), 108 calcis {Bacterium), 108 calcis (Pseudomonas), 108 calco-acetica {Pseudomonas), 146 calco-aceticus {Micrococcus), 255 calendulae {Bacterium), 133 calendulae {Phytommias) , 133 calendulae (Pseudomonas), 133 calf actor {Bacillus) , 740 calidolactis (Bacillus), 732 calidus {Bacillus), 732 California {Salmonella) , 505 californicus {Actinomyces) , 936 californicus (Streptomyces), 936 calligyra {Spirochaeta) , 1072 calligyrum (Treponema) , 1072 callistephi (Chlorogenus), 1146 callistephi var. attenuatiis {Chlorogenus), 1147 callistephi var. californicus {Chloroge- nus), 1147 caloritolerans (Clostridium), 797 caloritolerans {Pleciridium), 797 Calymmatobacterium, 14, 457 cameli {Actiiiomyces) , 915 cameli {Oospora), 915 cameli {Streptothrix) , 915 camelidae {Treponema) , 1076 campeneus {Micrococcus), 255 campestre (Marmor), 1202 campestre var. armoraciae {Bacterium) , 156 campestre var. galbinum {Marmor), 1202 campestre var. typicum {Marmor), 1202 campestris {Bacillus), 155 campestris {Bacterium) , 155 campestris {Phytomonas) , 155 campestris {Pseudomonas), 155 campestris (Xanthomonas), 155, 156, 160, 164, 178, 1134, 1136 campestris var. armoraciae {Phytomo- nas), 156, campestris var. armoraciae {Xanthomo- nas), 156, 164 canadensis {Bacillus) , 759 canadensis {Bacterium) , 759 canadiense {Clostridium), 819 1409 cana-capr INDEX OP NAMES OF GENERA AND SPECIES canalcnsis (Bacillus), 651 canaliculatus (Bacillus), 740 canalis (Bacterium) , 675 canalis (Microspira), 202 canalis capsulatus (Bacillus), 675 canalis capsulatus (Bacterium), 675 canalis parvum (Bacterium), 675 canalis parvus (Bacillus), 675 canariensis (Bacillus), 530 canastel (Salmonella) , 521 canceris (Bacidus), 740 cancrosi (Bacterium), 587 candicans (Achromobacter) , 423 candicans (Albococcus) , 255 candicans (Bacillus), 423 candicans (Bacterium) , 423 candicans (Coccus), 261 candicans (Micrococcus), 251, 252, 255, 257, 258, 259, 260, 262, 263, 264, 268, 269, 270, 273, 274, 275, 276, 277, 278, 279, 280, 281 candicans (Staphylococcus) , 255 Candida (Nocardia), 968 Candida (Sarcina), 290 Candida (Streptothrix) , 934, 970 candidus (Actinomyces) , 968 candidus (Bacterium), 401 candidus (Discomyces) , 968 candidus (Micrococcus), 239, 251, 252, 253, 254, 255, 256, 258, 264, 265, 267, 269, 270, 271, 272, 274, 275, 278, 279, 281, 282 candidus (Staphylococcus) , 239, 282 canescens (Albococcus), 255 canescens (Micrococcus), 255 canescens (Sarcina), 290, 291 canescens (Staphylococcus), 255 caneus (Lactobacillus), 363 canicida (Bacterium), 553 canicola (Leptospira), 1077, 1079 canidae (Treponema), 1076 canina (Palmula), 812 canina (Spirochaeta) , 1066 caninus (Acujormis), 812 caniperda (Bacillus), 740 canis (Actinomyces) , 915 canis (Asterococcus) , 1292 canis (Bacterium), 740 cants (Bartonella), 1104 cants (Cladothrix) , 915 cants (Corynethrix), 406 cants (Ehrlichia), 1096 cants (Ehrlichia) (Rickettsia), 1095 canis (Haemobartonella), 1104, 1105, 1106 cants (Hemophilus) , 587 cants (Nocardia), 915 cants (Oospora), 915 cants (Rickettsia), 1095 cants (Spirella), 217 cants (Spirochaeta), 1066 cants (Streptothrix), 915 canis (Tarpeia), 1272 canis familiaris (Pleuromyces) , 915 cants lupus (Grahamella) , 1110 caniseptica (Pasteurella) , 553 cannabinus (Bacillus), 813 cannae (Bacterium), 171 cannae (Phytomonas), 171 cannae (P seudomonas) , 171 cannae (Xanthomonas), 171 cantabridgensis (Ascococcus) , 250 canns (Bacillus), 651 canus (Micrococcus), 267 capillaceus (Bacillus), 740 capillorum (Micrococcus), 255 capillorum (Palmella), 255 capillorum (Palmellina), 255 capillorum (Zoogloea), 255 capillosa (Ristella), 577 capillosus (Bacillus), 577 capitovale (Clostridium), 795 capitovalis (Bacillus), 795 capitovalis (Plectridium) , 795 cappelletli (Streptococcus) , 317 caprae (Actinomyces), 899 caprae (Cladothrix) , 899 caprae (Discomyces) , 899 caprae (Nocardia), 899 caprae (Oospora), 899 caprae (Streptothrix) , 899 caprt (Bacillus), 714 capriformis (Micrococcus) , 255 caprinus (Streptococcus), 338 1410 INDEX OF NAMES OF GENERA AND SPECIES capr-caro capriseptica (Pasteurella) , 553 caprisepticus (Bacillus), 553 caprogenes (Bacterium) , 753 caprogenes foetidus (Bacillus), 753 caproicum (Clostridium), 820 capsaformans (Micrococcus), 255 c.apsici (Bacillus), 740 Capsularis, 33, 34, 577 capsulata (Klebsiella), 459 capsulata (Pseudomonas), 93, 146 capsulata (Rhodosphaera) , 865 capsulatum (Acetobacter), 189 capsulatum (Bacterium), 459 capsulatum (Rhodohacterium) , 863 capsulatum (Rhodonostoc) , 864 capsulatus (Bacillus), 459 capsulatus (Bacterium), 691 capsulatus (Diplococcus), 308 capsulatus (Mycococcus), 891 capsulatus (Rhodococcus) , 8, 865 capsulatus (Rhodopseudomonas), 864, 866 capsulatus (Rhodorrhagus) , 865 capsulatus (Streptococcus) , 308, 338 capsulatus aerogenes (Bacillus), 789, 790 capsulatus anaerohius (Bacillus), 790 capsulatus chinensis (Bacillus), 456 capsulatus gallinarum (Streptococcus), 338 capsulatus margarineus (Diplococcus), 662 capsulatus inucosus (Bacillus), 459 capsulatus mucosus (Bacterium), 459 capsulatus pyaemiae cuniculi (Bacillus) , 459 capsulatus septicus (Bacillus), 691 capsulatus septicus (Bacterium), 691 capsulatus septicus (Protetis), 691 carabiformis (Bacillus), 651 caraibica (Microspira) , 636 caraibicum (Photobacterium) , 636 carateum (Treponema), 1072 carbo (Micrococcus), 255 carbonei (Clostridium), 807 carbonei (Inflabilis) , 807 carbonis (Bacillus), 776 Carboxydomonas, 20, 31, 972 Cardiff (Salmonella), 512 cardn' (Fiferio), 203, 205 cardii-papiltosi (Cristispira) , 1056 cardii papillosi (Spirochaeta) , 1056 cardio-arthritidis, 320 cardiopyrogenes (Spirillum), 217 carduus (Coccus), 250 caricae (Marmor), 1201 carts (Bacillus), 816 carnea (Cladothrix) , 968 carnea (Nocardia), 968 carnea (Oospora), 968 carnea (Sarcina), 290 carnea (Streptothrix) , 968 carnegieana (Erwinia), 468 carneum (Bacterium) , 675 carneus (Actinomyces), 968 carneus (Bacillus), 675 carneus (Discomyces) , 968 carneus (Micrococcus) , 255 carneus (Streptococcus), 338 carneus halophilus (Tetracoccus), 284 carnicolor (Bacillus) , 675 carnicolor (Micrococcus), 255, 276 carniphilus (Bacillus), 740 carniphilus (Micrococcus), 255 carnis (Bacillus), 651, 810 carnis (Clostridium), 810 carnis (Plectridium) , 810 carnis (Streptococcus), 338 carnis foeiidum (Clostridium), 787 carnis saprogenes (Bacillus), 782 carnofoetidum (Clostridium), ISl carnosum (Bacterium), 675 carnosus (Actinomyces), 968 carnosus (Bacillus), 740 carocyanea (Pseudomonas), 146 carocyaneum (Bacterium), 146 carocyaneus (Bacillus), 146 carogeanui (Cohnistreptothrix) , 928 Carolina (Salmonella) , 531 carolinus (Bacillus), 531 carotae (Phytomonas) , 165 carotae (Pseudomonas) , 165 carotae (Xanthomonas), 165 carotarum (Bacillus), 714 carotarum (Bacterium), 714 1411 caro-cata INDEX OF NAMES OF GENERA AND SPECIES carotovora (Erwinia), 469, 470, 471, 474, 1129, 1134, 1135, 1136 carotovorum {Bacterium) , 469 carotovorum (Pectobacterium), 464, 469 carotovorus (Bacillus), 469 carougeaui (Actinomyces), 928 carougeaui (Discomyces), 928 carougeaui (Nocardia), 928 carougeaui (Streptothrix) , 928 carpanoi (Treponema), 1074 carpathiens (Bacillus), 361 Carphococcus, 235 Carpophthora, 1151 carrau (Salmonella), 528 carrosus (Streptococcus), 338 carteri (Borrelia), 1061, 1068 carteri (Spirillum), 1061 carteri (Spirochacta) , 1061 carteri (Spironema), 1061 carteri (Spiroschaudinnia) , 1061 carteri (Treponema), 953 Carteria, 925 Carterii, 925 cartharinensis (Micrococcus), 255 cartilagineum (Bacterium), 675 cartitagineus (Bacillus), 675 Caryococcus, 1121 Caryophanon, 1004 caryophyllacearum (Bacillus) , 639 caryophylli (Phytomonas) , 137 caryophylli (Pseudomonas), 136 raset (Actinomyces), 968 casei (Bacterium), 718 casez (Cellulomonas), 619 casei (Lactobacillus), 356, 357 casei (Micrococcus), 240 casez (Planococcus) , 281 casei (Plocamohacterium) , 695 casei (Propionihacterium) , 379 casei (Sarcina), 290 cosei (Streptohacterium) , 356, 361 casei (Streptococcus), 338 casei (Tetracoccus), 284 casei a (Bacillus), 356 casei 7 (Bacillus) , 358 casei 5 (Bacillus) , 360 casei e (Bacillus), 352 casei a (Bacterium), 356 casei « (Bacterium), 352 casei acido-proteolyticiis I (Micrococcus) , 240, 326 casei acido-protcolyticus II (Micrococ- cus), 240, 326 casej amari (Micrococcus), 326 casei amari edamicus (Micrococcus), 256 casei filans (Bacterium) , 354 casei limburgensis (Bacillus) , 612 casei limburgensis (Bacterium) , 612 casei liquefaciens (Micrococcus), 240, 326 casei liquefaciens (Tetracoccus), 240 casei proteolyticus I and II (Micrococcus) , 327 caseia (Cellulomonas), 619 caseicola (Bacterium), 676 caseinicum (Achromobacler) , 692 Caseobacterium, 8, 349 Caseobacterium e, 352 Caseococcus, 327 caseolytica (Sarcina), 291 caseolyticum (Bacterium), 371 caseolyticus (Bacillus) , 651 caseolyticus (Micrococcus), 240, 258, 259, 260, 263, 265, 266, 268, 273, 695 cassavac (Bacterium), 466 cassavae (Erwinia), 466 castaneae (Bacterium), 138 castaneae (Phytomonas), 138 castaneae (Pseudomonas), 138 castanicolum (Bacterium) , 640 castellanii (Castellanus), 542 castellanii (Micrococcus), 255 castellanii (Rhodococcus) , 255 castellanii (Shigella), 542 Castellanus, 535 castellum (Bacterium), 676 castigata (Cellulomonas), 615 castigatum (Bacterium), 615 castra (Vibrio), 203 catarrhalis (Actinomyces), 922 catarrhalis (Bacillus), 590 catarrhalis (Discomyces), 922 catarrhalis (Micrococcus), 298 catarrhalis (Neisseria), 298, 299, 301 catarrhalis (Oospora), 922 1412 INDEX OF NAMES OF GENERA AND SPECIES cata-cell catarrhalis (Pseudotnonas), 146 Catenabacterium, 33, 34, 368 catenajorme {Catenahacterium), 368 catenaformis {Bacteroides) , 368 catenatus {ThermobaciUus) , 731 catenula (Bacillus), 676, 823 catenula (Bacterium), 676 catenula (Cornilia), 823 catenula (Tyrothrix), 823 catemdatus (Bacillus), 740 catenulatus (Chondromyces), 1039 cathetus (Bacillus), 651 cati (Actinomyces) , 969 cati (Discomyces) , 969 cattleyae (Bacterium) , 759 caucasica (Dispora), 351 caucasica (Pacinia), 351 caucasicum (Bacterium) , 351 caucasicwn (Betabacterium), 351, 358 caucasicus (Bacillus), 351, 358 caucasicus (Lactobacillus), 351 caucasicus (Streptococcus), 338 caudata (Pseudomonas), 174, 405 caudatum (Flavobacterium) , 174 caudatus (Bacillus), 174, 405 caudatus (Bacterium), 174 caudatus (Coccus), 250 Caulobacter, 35, 832 cavarum pericarditis (Bacterium) , 554 cavalum (Bacterium) , 676 cavernae (Bacterium), 676 cavernae minutissimus (Bacillus), 676 caviae (Actinomyces), 912 caviae (Bacillus), 651 caviae (Bacteroides), 574, 580 caviae (Bartonella) , 1108 caviae (CristispireUa), 1057 caviae (Haemobartonella) , 1108 caviae (Klebsiella), 548 caviae (Nocardia), 912 caviae (Pasteurella), 553, 651 caviae (Pseudomonas) , 146 caviae (Spherophorus) , 574, 580 caviae (Spirochaeta) , 1071 caviae (Spironema), 1070 caviae (Spiroschaudinnia) , 1070 caviae (Streptobacillus) , 574 caviae fortuitum (Bacterium), 676 caviae fortuitus (Bacillus), 676 cavicida (Bacillus), 445 cavicida (Bacterium) , 445 cavicida (Escherichia), 445 cavicida havaniensis (Bacillus), 676 cavicida havaniensis (Bacterium) , 676 caviseptica (Pasteurella), 553 cavisepticum (Bacteritim) , 553 cazaubon (Bacterium) , 759 cazaubon I and // (Bacterium), 759 celebense (Bacterium), 169 celebensis (Phytom,onas), 169 celebensis (Pseudomonas), 169 celebensis (Xanthomonas), 169 celer (Phagus), 1142 cellaris (Leucocystis) , 255 cellaris (Micrococcus) , 255 cellasea (Cellulomonas), 618 cellaseus (Bacillus), 618 Cellfalcicula, 211 cellobioparus (Clostridium) , 820 cellulicola (Schinzia), 224 Cellulobacillus, 8, 705, 763 cellulolyticum (Plectridium), 823 Cellulomonas, 20, 32, 615, 616 cellulomonas (Proteus) var. Proteus biazo- teus, 617 cellulomonas (Proteus) var. Proteus cas- taneus, 622 cellulomonas (Proteus) var. Proteus ros- sicus, 622 cellulomonas (Proteus) var. Proteus udus, 614 cellulosae (Actinomyces) , 938 cellulosae (Clostridium), 820 cellulosae (Streptomyces), 938 cellulosae dissolvens (Bacillus), 809 cellulosae dissolvens (Caduceus), 809 cellulosae hydrogenicus (Caduceus), 809 cellulosae hydrogenicus var. cellulosae methanicus (Caduceus), 810 cellulosae methanicus (Caduceus), 810 cellulosam (Bacterium) , 808 cellulosam fermentans (Bacillus), 808 cellulosis (Bacterium) , 809 cellulosolvens (Caduceus), 809 1413 cell-ceyl INDEX OF NAMES OP GENERA AND SPECIES cellulosolvens (Clostridium), 809 cellulosum (Angiococcus), 1048 cellulosum (Polyangium), 1027, 1028 cellulosum (Sorangium), 1022 cellulosum fcrmentans (Terminosporus) , 808 cellulosum var. ferrugineutti (Polyan- gium), 1010, 1028, 1029 cellulosum var. fulvum (Polyangium), 1029 cellulosum var. fuscum (Polyangium), 1010, 1028 cellulosum var. luteum (Polyangium), 1029 Cellvibrio, 209 Cenomesia, 847 centrale (Anaplasina), 1100 centralis (Bacillus), 651 centricum (Bacterium) , 676 centrijugans (Bacil us), 101, 697 cenlrifugans (Bacterium), 697 centrijugans (Pseudomonas), 101, 697 centropunctatum (Achromobacter), 423 centropunctatus (Bacillus), 423 centropunclalus (Bacterium), 423 centropunctatus (Micrococcus), 255 centrosporogenes (Bacillus), 787, 825 centrosporogenes (Clostridium), 787 Centrosporus, 20, 22 centrosporus (Bacillus), 725, 744 cepae (Bacillus), 740 cepae (Marmor), 1184 cephaloideus (Bacillus), 816 cepivora (Phytomonas), 470 cepivorum (Bacterium), 470 cepivorus (Aplanobacter) , 470 cepivorus (Bacillus), 470 ceramicola (Bacterium), 626 ceramicola (Flavohacterium), 626 cerasi (Bacterium), 120 cerasi (Marmor), 1197 cerasi (Phytomonas), 120 cerasi var. prunicola (Pseudomonas), 120 cerasi wraggi (Bacterium), 153 cerasi wraggi (Phytomonas), 153 cerasinus (Micrococcus), 255, 256, 338 cerasinus (Streptococcus), 338 cerasinus lactis (Micrococcus), 255 cerasinus siccus (Micrococcus), 338 cerasus (Bacillus), 120 cerasMS (Pseudomonas), 120 cereale (Lactobacillus), 355 cereale (Thermobacterium), 355 cerealia (Pseudomonas) , 740 cerealium (Bacillus), 740 cerealium (Bacterium) , 740 cerebralis (Corynebacterium), 403 cerebriformis (Actinomyces), 969 cerebriformis (Chondrococcus), 1046 cerebriformis (Myxococcus) , 1046 cerebriformis (Nocardia) , 969 cerebriformis (Streptothrix) , 969 cereus (Actinomyces) , 969 cereus (Bacillus), 708, 715, 716, 717, 718, 719, 725, 1138 cereus (Micrococcus), 256 cereus (Staphylococcus) , 251 cereus albus (Micrococcus) , 251 cereus albus (Staphylococcus), 251 cereus aureus (Micrococcus), 256 cereus aureus (Staphylococcus) , 256 cereus Jlavus (Micrococcus), 256 cereus flavus (Staphylococcus) , 256 cereus var. Jluorescens (Bacillus), 716 cereus var. mycoides (Bacillus), 718 cereus var. siamensis (Bacillus) , 716 cerevisiae (Flavobacterium) , 176 cerevisiae (Lactobacterium) , 363 cerevisiae (Merismopedia) , 249 cerevisiae (Micrococcus), 249 cerevisiae (Pediococcus), 249 cerevisiae (Pseudomonas), 176 cerevisiae (Sarcina), 249 cerinum (Bacterium), 676 cerinus (Gluconoacetobacter) , 694 cerinus (Micrococcus), 256 cerro (Bacterium), 529 cerro (Salmonella), 529 ceruminis (Bacillus), 403 ceruminis (Corynebacterium) , 403 cervina (Sarcina), 291 ceylonensis (Shigella), 542, 543 1414 INDEX OF NAMES OF GENERA AND SPECIES ceyl-chro ceylonensis A (Bacillus), 540 ceylonensis B (Bacillus) , 542 ceylonensis B (Lankoides) , 542 chalcea (Actinomyces), 978 chalcea (Micromonospora) , 959, 978 chalcea (Nocardia), 978 chalcea (Streptothrix) , 978 chalmersi (Actinomyces), 975 chalmersi (Nocardia), 975 chamae (Cristispira), 1056 chamae (Spirochaeia) , 1056 Charon, 1265 charrini (Bacillus), 651 charrini (Streptococcus), 338 chauvaei (Bacillus), 776 chauvaei (Clostridium) , 776 chauvei (Clostridium) , 776 Chauvoea, 20, 22, 763 chauvoei (Bacillus), 776 chauvoei (Bacterium) , 776 chauvoei (Buty r ibacillus) ,77Q chauvoei (Clostridium) , 776 chelonei (Mycobacterium), 886 chersonesia (Micrococcus), 256 Chester (Salmonella), 504 chinense (Aerobacter) , 456 chinense (Bacterium), 456 chinicus (Micrococcus) , 256 chironomi (Bacterium) ,^"^5 chironojni (Photobacterium) , 635 chitinochroma (Bacterium), 632 chitinophilum (Bacterium), 631 chitinovorus (Bacillus), 632 Chlamydothrix, 12, 18, 984 Chlamydozoon, 1114 chlorina (Pelogloea), 870 chlorina (Pseudomonas), 95 chlorinum (Bacterium), 95, 654, 676 chlorinus (Bacillus), 95, 651 chlorinus (Micrococcus) , 256 Chlorobacterium, 693, 872 Chlorobium, 29, 30, 869 Chlorochromatium, 873 Chlorogenus, 1146 Chloronium, 873 Chloronostoc, 869 ChloTophaena (Pseudomonas) , 146 Chloropseudomonas, 870 chlororaphis (Bacillus) , 93 chlororaphis (Pseudomonas), 93 chlorum (Flavobacterium) , 440 chocolatum (Chromobacterium) , 693 cholerae (Babillus), 194 cholerae (Bacterium) , 547 cholerae (Phagus), 1142 cholerae (Vibrio), 194, 195, 1142 cholerae anatum (Bacillus), 552 choleraeasiaticae (Pacinia), 193 cholerae asiaticae (Spirillum), 193 cholerae asiaticae (Vibrio), 194 cholerae -caviae (Bacillus), 502 cholerae columbarum (Bacillus), 552 cholerae columbarum (Bacterium) , 552 cholerae gallinarum (Bacillus), 547 cholerae gallinarum (Bacterium) , 547 cholerae gallinarum (Micrococcus) , 547 cholerae gallinarum (Octopsis), 547 cholerae gallinarum (Pasteurella), 547 cholerae-suis (Bacillus), 508 cholerae-suis (Bacterium), 508 cholerae-suis (Bacterium) (Salmonella), 508 choleraesuis (Salmonella), 493, 494, 496, 508,509 cholerae suis var. kunzendorj (Salmon- ella), 509 510 cholerae suum (Bacillus), 508 cholerae suum (Bacterium), 508 choleroides (Bacillus), 198 choleroides (Bacterium), 702 choleroides (Microspira), 198, 203 choleroides a and |3 (Fi6/io), 203, 702 cholesterolicum (Mycobacterium) , 890 chologenes (Bacillus), 676 chologenes (Bacterium), 676 chondri (Streptothrix) , 976 Chondrococcus, 1009, 1044 Chondromyces, 14, 17, 20, 24, 26. 1036 choukevitchi (Bacillus), 612 christiei (Bacterium), 759 christophersoni (Actinomyces) , 975 christophersoni (Nocardia), 975 Chromatium, 16, 23, 25, 29, 30, 846, 852, 853, 856, 857, 859 1415 chro-citr INDEX OF NAMES OF GENERA AND SPECIES chromidrogenus citreus (Micrococcus) , 256 chromidrogenus ruber (Micrococcus), 256 chromoaromaticus (Bacillus), 657 chromo-aromaticus (Bacterium), 657 Chromobacterium, 20, 32, 37, 223, 231, 694 chromoflavus (Micrococcus), 256 chromogenes (Cladothrix) , 969 chromogenes (Clostridium), 805 chromogenes (Oospora), 969 chromogenes ^ alba, (Actinomyces) 934 chromogenus (Actinomyces), 934, 940, 969, 970, 972 chromogenus 205 (Actinomyces) , 941 chroococcum (Azotobacter), 219 chroococcus (Bacillus) , 219 Chroostipes, 872 chrysanthemoides (Vibrio), 203 chryseum (Bacterium) , 676 chryseus (Bacillus) , 672 chryseus (Micrococcus), 256 chrysogloea (Bacterium) , 676 chrysogloia (Bacillus), 676 chylogena (Eberthella) , 533 chylogenes (Bacillus), 450, 533 chyluriae (Bacillus), 651 chymogenes (Bacterium), ^52 dehor ii (Bacterium), 133 cichorii (Phytomonas), 133 cichorii (Pseudomonas), 125, 133 ciliatus (Lactobacillus) , 363 Cillobacterium, 33, 34, 369 cincinnatus (Bacillus), 813 cinctus (Bacillus) , 740 cinerea (Neisseria), 299, 301 cinereo-niger (Actinomyces) , 969 cinereonigeraromaticus (Actinomyces), 969 cinereo-nigra (Nocardia), 969 cinereus (Micrococcus), 301 cinereus (Streptococcus), 338 cinereus niger aromaticus (Actinomyces), 969 cineronigra aromatica (Streptothrix), 969 finnabareus (Micrococcus), 244, 255; 256, 257, 274, 275 cinnabareus (Rhodococcus), 244 cinnabar iyius (Micrococcus) , 256 circulans (Bacillus), 722, 728, 737, 741, circulans (Bacterium), 722 circularis major (Bacillus), 580 circularis minor (Bacillus), 362 cirrhijormis (Micrococcus), 256 cirrhosus (Chondrococcus), 1045 cirrhosus (Myxococcus), 1045 cirroflagellosus (Bacillus), 741 cissicola (Aplanobacter) , 134 cissicola (Pseudomonas), 134 CiiitJir, 1209 citrarefaciens (Bacterium), 119 citrarefaciens (Pseudomonas), 119 citrea (Nocardia), 908, 975 citrea (Sarcina), 288, 291 citrea (Streptothrix), 969 citrea conjunctivae (Sarcina), 291 citreum (Bacterium) , 651, 676, 687 citreum (Semiclostridium) , 762 citreus (Actinomyces) , 946, 969 citreus (Ascobacillus) , 651 citreus (Bacillus), 651, 676 citreus (Enterococcus) , 336 citreus (Micrococcus), 239, 242, 254, 256 257, 261, 265, 268, 278, 280, 338 citreus I (Micrococcus) , 256 citreus II (Micrococcus) , 256, 275 citreus (M ycococcus) , 891 citreus (Planococcus) , 288 citreus (Proactinomyces), 908 citreus (Staphylococcus) , 242 citreus (Streptococcus) , 338 citreus (Streptomyces), 946 citreus baregensis (Microbacillus) , 690 citreus cadaversis (Bacillus), 687 citreus cadaveris (Bacterium), 687 citreus conglomeratus (Diplococcus) , 239 citreus conglomeratus (Merismopedia) , 239 citreus congtomeratus (Micrococcus), 239 citreus duodenalis (Staphylococcus), 701 citreus granulatus (Micrococcus) , 256, 277 citreus lactis (Micrococcus), 257 citreus liquefaciens (Diplococcus) , 257 citreus liquefaciens (Micrococcus), 257 citreus rigensis (Micrococcus), 257 cziri (Bacillus), 156 1416 INDEX OF NAMES OF GENERA AND SPECIES citr-cocc citri {Bacterium) , 156 citri (Phagus), 1135 citri (Phyiomonas) , 156 citri (Pseudomonas) , 156 citri (Xanthomonas), 156, 1129, 1134, 1135, 1136 citri deliciosae {Bacterium), 17S citricus {Bacillus), 651 citrimaculans {Bacillus), 475 citrimaculans {Bacteriu77i), 475 citrimaculans (Erwinia), 475 citrina {Sarcina), 291 citrinus {Bacillus) , 651 citrinus {Micrococcus), 257 citriputeale {Bacterium), 119, 120, citriputealis {Phytomonas) , 120 citriputealis {Pseudoinonas), 120 Citrobacter, 448 citrocremeus {Actinomyces), 915 citrophilum (Achromobacter), 610 citrophilum {Urobacterium), 610, 691 citrophilus {Streptococcus), 339 citrovorum (Leuconostoc), 347 citrovorus {Streptococcus) , 347 citrovorus-paracitrovorus {Streptococcus), 348 Cladascus, 12, 13 cladogenes {Bacillus), 651 cladoi {Bacillus), 741, 75S Cladolhrix, 6, 17, 18, 917, 980, 982, 1 121 claibornei {Salmonella) , 518 clathratiforme {Aphanothecc) , 871 clathratiforme (Pelodictyon), 871, 872 Clathrochloris, 872 Clathrocystis, 6, 847 clavatus {Bacillus), 400, 813 clavatus {Myxococcus), 1045 clavifer {Actinomyces) , 969 clavifolium (Aureogenus), 1157 claviforme {Bacterium), 823 claviformis {Bacillus), 651, 823 claviformis {Micrococcus), 257 claviformis {Pacinia), 823 claviformis {Tyrothrix), 823 demo {Pseudomonas), 694 cleoni {Bacillus), 652 Cloaca, 10 cZoaca {Microspira) , 202, 203, 206 cloacae {Actinomyces), 969 cloacae (Aerobarter), 455, 456, 457, 460, 670, 692 cloacae {Bacillus), 455, 457 cloacae {Bacterium), 455 cloacae {Cloaca), 455 Clonothrix, 12, 17, 19, 26, 35, 983 clonotricoides {Mycothrix), 983 closteroides {Bacillus), 722, 741 clostridiiformis (Bacterium), 576, 577 clostridiiformis {Ristella), 576 clostridiiformis mobilis {Znberella), 577 clostridioides {Bacillus), 720 Clostridium, 11, 22, 27, 30, 31, 33, 42, 43, 76, 216, 367, 763 Clostrillium, 7, 705 Clostrinium, 7, 705 coadunata (Pseudomonas), 101, 697 coadunatum {Achromobacter) , 101 coadunatus {Bacillus), 101 coadunatus {Bacterium), 697 coagulans (Bacillus), 531, 713 coagulans (Bacteroides), 567, 577 coagidans {Balkanella) , 531 coagulans {Clostridium), 782 coagulans {Pasteurella), 567 coagulans {Salmonella), 531 cobayae {Bacillus), 714 cobayae {Borrelia), 1066 cobayae {Spirochaeta) , 1066 cobayae {Treponema), 1066 coccacea {Pseudomonas) , 146 cocciforme {Bacterium), 403, 693 cocciformis {Brucella), 693 coccineum {Thiospirillum) , 856, 859 coccineus {Bacillus), 652, 741 coccineus {Micrococcus) , 257 Coccobacillary bodies (Nelson), 1294 Coccobacillus, 546, 1291 Coccobacterium, 479 coccoides (Eperythrozoon), 1112, 1113 coccoides (Nitrosocystis), 72, 73 coccoideum {Achromobacter) , 423 coccoideum {Bacterium), 423 coccoideus {Bacillus) , 710 Coccomonas, 11 1417 cocc-coli INDEX OF NAMES OF GENERA AND SPECIES Coccothrix, 876 cochleariuin (Clostridium), 794, 799 cochlearinm (Plcctriduim), 794 cochlear ills (Bacillus), 794 cochlearius [Flemingillus) , 794 cocoide (Bartonella) , 1101 cocci (Bacillus), 416 coeliaca (Xocardia), 899, 908, 913 coeliacum (Flavohacterium) , 906 coeliacum (Mycobacterium) , 392, 906 coeliacus (Proactinomyces), 906 coelicolor (Actinomyces), 935 coelicolor (Bacillus), 401 coelicolor (Bacterium), 401 coelicolor (Nocardia), 935 coelicolor (Streptomyces), 935 coelicolor (Streptothrix) , 935 coeln (Salmonella), 503 coelorhynchus (Coccobacillus), 636 coenohios (Pseudomonas) , 697 cocrulea (Micromonospora), 980 coerulea (Pseudomo7ias), 234 coeruleO'Viridis (Bacillus), 652 coeruleum (Chromobacterium) , 234 coeruleus (Bacilhis), 234, 652 coeruleus (Bacterium), 234 coffeicola (Bacillus), 639 cohaerea (Pseudomonas), 146, 697 cohaerens (Bacillus), 146, 632, 697, 718 cohaerens (Chromobacterium) , 234 cohaereus (Bacterium), 697 Cohnia, 6, 847 cohnii (Spirillum), 1062 cohnii (Spirochaeta) , 1062 cohnii (Spirochaete), 1054 Cohnistreptothrix, 925, 929 Coleomitus, 1003 Coleonema, 1003 Colesiota, 1119 corj (Acrobacicr), 445 coZi (Bacillus), ii^ coli (Bacterium), 445, 460 coh' (Colobaclrum), 445 coli (Escherichia), 3, 444, 445, 447, 448, 449, 450, 451, 454, 455, 460, 489, 595, 694, 1131, 1132, 1133, 1134. 1296 coli (Phagus), 1133 coli (Pseudomonas) , 146 coil (Streptococcus) , 339 co?^ alcaligenes (Bacterium), 452 co^l anaerogenes (Bacillus) , 533 coZi anaerogenes (Bacterium), 533, 542 colianaerogenes (Castellanus), 533 coZi anindolicum (Bacterium), 452 coZ? apium (Bacterium) , 676 coZz brevis (Micrococcus), 257 coiz brevis (Streptococcus) , 339 coh' citrovorum (Bacterium) , 448 coZi color abilis (Bacillus) , 612 coZz colorabilis (Bacterium), 612 coZi commune (Bacterium), 445 coh' communior (Bacillus), 447 coli-communior (Bacterium), 447 coZz communis (Bacillus), 445, 632, 697 co/i communis (Pseudomonas), 697 coiz communis verus (Bacillus), 445 coil dysentericum (Bacillus), 543 coli flavum (Bacterium) , 445 coh" gracilis (Streptococcus), 326 coil immobilis (Bacillus), 450 co/i immobilis (Bacterium), 450 coZi imperfectum (Bacterium, 452 coh' mobilis (Bacillus), 148 coZt mobilis (Bacterium) , 699 coh" mutabile (Bacterium), 451 coiz mutabile (Escherichia) , 451 coil mutabilis (Bacillus) , 450 coh' mutabilis (Escherichia), 451 coZi similis (Bacillus) , 676 co/z similis (Bacterium), 676 coiz var. acidilactici (Bacterium), 447 coZi var. acidilactici (Escherichia), 447 coh' var. communior (Bacterium), 448 coZi var. communior (Escherichia), 447 coii var. immobilis (Bacterium) , 450 coZi var. luteoliquefaciens (Bacillus), 490 coh' var. luteoliquefaciens (Bacterium), 490 co/z var. neapolitana (Escherichia), 447 coii var. neapolitanum (Bacterium), 447 coil var. paragruenthali (Bacterium), 451 coil wrws (Bacillus), 445 Colibacter, 694 Colibacterium, 11, 444 1418 INDEX OF NAMES OF GENERA AND SPECIES coli-conj colicogenes (Bacillus), 813 coliforme (Paracolobactrum), 460, 491 Colloides, 595 colloides (Pseudomonasj, 146 colloideum (Bacterium), 676 Colobactrum, 444, 453 colofoetida (Escherichia), 452 colofoetidus (Bacillus) , 452 coloides (Escherichia), 452 coloides var. A (Bacillus), 452 coloides var. B (Bacillus), 452 colomatii (Bacterium), 759 colorabilis (Bacillus) , 612 colorans (Bacillus), 741 coloratum (Propionibacterium) , 379 colossus (Spirillum), 217 colotropicalis (Bacillus) , 452 colotropicalis (Escherichia), 452 columbarwn (Bacillus), 652 columbarum (Bacterium), 401, 552 columbense (Bacterium), 531 columbensis (Bacillus), 531 columbensis (M organella), 531 columbensis (Salmonella) , 531, 532 columnaris (Bacillus), 1047 columnaris (Chondrococcus), 1047 columnaris (Cytophaga), 1047 colurnae (Phytomonas), 139 colurnae (Pseudomonas), 139 comandoni (Spirochaeta) , 1066 romandoni (Treponema), 1066 comes (Bacterium) , Q7Q comesii (Bacillus), 741 comma (Bacillus) , 194 comma (Microspira) , 194 comma (Vibrio), 193, 196, 198, 199, 202, 203, 204, 205, 206, 1142, 1143 commensalis (Diplococcus), 257 commensalis (Micrococcus), 257 commune (Colibacter), 694 commune (Corynebacterium), 403 commune (Semiclostridium) , 762 communior (Bacillus), 447 communior (Bacterium), 447 communior (Escherichia) , 448 communior var. coscoroba (Bacillus), 453 communis (Bacillus), 445, 681 communis (Siderococcus) , 835 communis lactis (Micrococcus), 257 commutabilis (Phagus), 1136 commutatus (Micrococcus), 257 compactum (Bacterium), 676 compactus (Bacillus), 676 compositum (Polyangium), 1023 compositum (Sorangium), 1008, 1022, 1023 C07icentricum (Bacterium), 676 concentricum (Spirillum), 217 concentricus (Micrococcus), 257 concitata (Cellulomonas), 619 concitatus (Bacillus), 619 concoctans (Bacillus), 741 concomitans (Bacillus), 230 concretivorus (Thiobacillus), SI confervarum (Siderocystis) , 834 confervarum (Sideromonas), 834 confluens (Micrococcus), 257 conglomeratum (Lactobacterium) , 363 conglomeratus (Micrococcus), 239, 252, 253, 256, 257, 258, 261, 262, 265, 270^ 271, 280 conglomeratus (Streptococcus), 315 congolensis (Actinomyces) , 918 congolensis (Discomyces) , 918 congregata (Sporocytophaga), 1049, 1050 Conidiothrix, 984, 995 conjac (Bacterium), 171 conjac (Phytomonas), 171 conjac (Pseudomonas), 171 conjaci (Xanthomonas), 171 conjunctivae (Chlamydozoon), 1119 conjunctivae (Colesiota), 1119, 1120 conjunctivae (Micrococcus), 257 conjunctivae (Rickettsia), 1096, 1119 conjunctivae (Sarcina), 291 conjunctivae bonis (Rickettsia), 1096, 1120 conjunctivae-galli (Colesiota), 1120 conjunctivae galli (Rickettsia), 1096, 1120 conjunctividis (Hemophilus), 585 conjunctividis (Micrococcus) , 257 conjunctivitidis (Bacillus), 677 conjunctivitidis (Bacterium), 676, 677 conjunctivitidis subtiliformis (Bacillus), 741 1419 conj-cory INDEX OF NAMES OF GENERA AND SPECIES conjunctivitis {Bacterium), 590 connii (Achromobacter) , 423 connii {Bacterium), 423 conoideus {Micrococcus), 257 conori {Dermacentroxenus) , 1088 conorii (Rickettsia), 1087, 1088, 1092 conradi {Pseudomonas), 146 consolidus {Bacillus) , 741 constans (Marmor), 1167 conslellatum {Bifidibacteri um) , 369 constellatus {Bacillus), 369 constellatus (Diplococcus), 310 constrictus {Bacillus), 652 constrictus {Bacterium), 652 contextus {Bacillus), 741 continuosus {Streptococcus), 339 coiitumax (Phagus), 1136 convexa {Pasteurella), 571 convexa (Pseudomonas), 96, 697. convexus (Bacteroides), 571, 577 convoluta {Nocardia), 919 convoluta {Oospora), 919 convoluturn {Mycobacterium), 919 convolutus {Actinomyces), 919 convolutus {Bacillus), 652 convollus {Bacterium) , 652 convolutus {Discomyces) , 919 coprocinus {Bacillus), 753 coprogenes, {Bacillus) 813 coprogenes foetidus {Bacillus), 652, 813 coprogenes parvus {Bacillus), 544 coprogenese parvus {Bacterium), 544 coproliticus {Thiobacillus), 80 coprophila {Microspira), 202, 206 coprophilum {Bacterium) , 819 coprophilum {Spirillum), 206 coprophilus {Bacillus), 819 coralinus {Rhodococcus) , 258 corallina (Nocardia), 897, 902, 903, 904 corallina {Pseudomonas) , 697, 1016 corallina {Serratia), 644, 902 corallinum {Clostridium) , 820 corallinus {Micrococcus) , 258 corallinus (Proactinomyces) , 902 corallinus {Streptohrix) , 902 coralloides (Chondroooocus), 1006, 1045, 1046 coralloides {Micrococcus), 258 coralloides {Myxococcus), 1045 coralloides var. clavatus {Chondrococcus) , 1045 coralloides var. polycystus {Chondrococ- cus) 1045 Cerium, 1203 corneola (Gallionella), 832 Cornilia, 705, 763 cornutum {Bifidibacterium) , 369 cornutus {Bacillus) , 369 cornutus {Bacteroides) , 369 coronafaciens {Bacterium), 116 coronafaciens {Phytomonas) , 116 coronafaciens (Pseudomonas), 113, 116 coronafaciens var. atropurpurea {Phy- tomonas), 116 coronafaciens var. atropurpurea {Pseu- domonas), 116 coronafaciens var. atropurpureum {Bac- terium), 116 coronata {Microspira), 636 coronata {Siderocapsa), 834 coronatum {Photobacterium) , 636 coronatus {Bacillus), 652 coronatus {Micrococcus), 339 coronatus {Streptococcus), 339 coroniformis {Actinomyces), 969 corrugatus {Bacillus), 741 corrugatus {Micrococcus), 258 corruleo-viride (Bacterium), 652 corruscans {Bacillus), 741 corticate {Bacterium) , 677 corticalis {Bacillus), 677 com {Bacillus) , 652 corylii {Bacterium) , 640 corylina {Phytomonas), 156 corylina (Xanthomonas), 156 Corynebacterium, 7, 17, 18, 21, 22, 23, 27, 30, 35, 37, 38, 42, 381, 382, 391, 396, 400, 401, 403, 404, 405, 407, 435, 612, 615, 633, 866, 927 Corynemonas, 8, 381 Corynethrix, 381 , 407 Corynobacterium, 12, 13, 19, 28, 381 coryzae {Diplococcus), 258 coryzae {Micrococcus) , 258 1420 INDEX OF NAMES OF GENERA AND SPECIES cory-cten coryzae contagiosae equon/m (Streptococ- cus), 317 coryzae segmeniosus [Bacillus), 406 coscoroba (Bacillus), 453, 557, 552 coscoroba (Escherichia), 453 coscorobae (Bacterium), 453 costatus (Bacillus), 741 costicolus (Vibrio), 702 costicolus var. liquefaciens (Vibrio), 702 cotti (Microspironenia), 1074 cotti (Treponema), 1074 couchi (Grahamella) , 1110 courmontii (Bacillus) , 652 couvyi (Leptospira), 1078 couvyi (Spirochaeta) , 1078 Cowdria, 1094, 1097 Coxiella, 1092 crassa (Klebsiella), 459 crassa (Leptothrix) , 985 crassa (Simonsiella) , 1004 crasse (Caryophanon), 1004 crassum (Bacterium) , 459 crassum (Plocamobacterium) , 400 crassum (Spirillum), 203, 217 crassum (Thiospirillum) , 851 crassMs (Bacillus), 362, 400, 652 crassus (Diplococcus) , 297, 301 crassus (Micrococcus) , 301 crassus (Ti;6ri'o), 203 crassus aromaticus (Bacillus), 146 crassus pyogenes (Bacillus), 652 crassus pyogenes bovis (Bacillus), 652 crassus sputi genus (Bacillus), 459 crassMS var. Z) (Fi6no), 203 crasieri (Vibrio), 204 craterifer (Actinomyces), 969 cremoides (Bacterium) , 403 cremoides (Corynebacteriurn) , 403 cremoides (Micrococcus), 253, 258 cremoides albus (Micrococcus), 258 cremoides aureus (Micrococcus), 253 cremoris (Bacillus), 709 rremoris (Streptococcus), 324, 325, 339, 340, 1138, 1139 cremorisviscosi (Micrococcus) , 258 cremoris-viscosi (Staphylococcus), 258 crenatum (Bacterium) , 677 crenatus, Thiobacillus, 81 Crenothrix, 6, 12, 17, 18, 19, 23, 26, 987 crepusculum (Micrococcus), 258 crepusculum (Monas), 258 cresologenes (Bacillus) , 813 cresologenes (Clostridium) , 813 cretacea (Oospora) , 969 cretaceus (Actinomyces), 969 cretaceus (Micrococcus), 258 cretus (Caryococcus), 1121 criceti domestici (Grahamella) , 1110 cricetuli (Grahamella), 1110 crinatum (Bacterium), 741 crinitus (Bacillus), 652, 741 cristalliferum (Bacterium) , 677 cristallino violaceum (Bacterium), 234 cristatus (Arthromitus), 1003 cristatus (Micrococcus) , 258 Cristispira, 12, 19, 20, 26, 28, 42, 1055, 1056. Cristispirella, 1069 crocatus (Chondromyces), 1006, 1036, 1038 crocea (Cytophaga) , 1016 crocz (Bacillus), 474 croci (Erwinia) , 474 crocidurae (Spirochaeta), 1066 crocidurae (Treponema), 1066 cromogena (Streptotrix), 969 crouposa (Klebsiella), 458 cruciferarum (Marmor), 1176 cruciformis (Micrococcus), 258 cruciviae (Achromobacter) , 103 cruciviae (Pseudomonas), 103 cruentus (Chondrococcus) , 1042 cruentus (Myxococcus), 1042 cruoris (Actinomyces) , 975 cruoris (Discomyces) , 975 cruoris (Nocardia), 975 cruoris (Oospora) , 975 crystalloides (Bacillus), 741 crystaloides (Bacterium) , 741 crystallophagum (Mycobacterium) , 897 crystallophagus (Actinomyces), 898 crystallophagus (Proactinomyces) , 898 ctenocephali (Rickettsia), 1096 ctenocephali (Spirochaeta), 1066 1421 cten-cuti INDEX OF NAMES OF GENERA AND SPECIES ctenocephali (Treponema), 1066 cubana (Salmonella), 527 cubensis (Bacillus), 741 cubensis (Spirochaeta) , 1066 cubonianum (Bacterium), 135, 693 cubonianus (Bacillus), 135, 652 cuculi (Bacillus), 403 cuculi (Corynebacterium) , 403 cuculliferum (Chromatium), 858 cucumeris (Lactobacillus) , 356 cucumeris (Marmor), 1155, 1173 cucumeris fermentati (Bacillus), 356 cucumeris jermentati (Ulvina), 695 cucumeris var. commelinae (Marmor), 1174 cucumeris var. judicis (Marmor), 1173, 1174 cucumeris var. lilii (Marmor), 1174 cucumeris var. phaseoli (Marmor), 1174 cucumeris var. upsilon (Marmor), 1172 cucumen's var. vignae (Marmor), 1174 cucumeris var. vulgare (Marmor), 1174 cucumis (Vibrio), 204 cucurbitae (Bacterium), 157 cucurbitae (Phytomonas) , 157 cucurbitae (Pseudomoiias) , 157 cucurbitae (Xanthomonas), 157 cuenoti (Bacillus), 652 culicis (Entomospira) , 1066 culicis (Rickettsia) , 1096 culicis (Spirillum) , 1066 culicis (Spirochaeta), 1066 culicis (Spironema), 1066 culicis (Spiroschaudinnia) , 1066 culicis (Treponema), 1066 cumini (Phytomonas), 121 cumini (Pseudomonas), 121 cumulatus (Micrococcus) , 258, 278 cumulatus tenuis (Micrococcus) , 278 cumulus yninor (Coccus), 694 cuneatum (Bacterium) , 776 cuneatus (Bacillus) , 813 cuneatus (Vibrio), 199, 203, 205 cuniculi (Actinomyces) , 578, 910, 928 cuniculi (Bacillus), 652 cuniculi (Bacterium) , 402, 552 cuniculi (Cladothrix) , 578, 928 cuniculi (Clostridium) , 820 cuniculi (Cohnistreptothrix) , 928 cuniculi (Corynebacterium) , 403 cuniculi (Hemophilus), 589 cuniculi (Klebsiella) , 459 cuniculi (Listerella) , 409 cuniculi (Nocardia), 910, 928 cuniculi (Noguchia) , 594 cuniculi (Oospora), 928 cuniculi (Pasteurella) , 547 cuniculi (Spirochaeta), 1073 cuniculi (Streptococcus) , 339 cuniculi (Streptothrix) , 578, 928 cuniculi (Treponema), 1073, 1076 cuniculi pneumonicum (Bacterium) , 552 cuniculi pneumonicus (Bacillus), 552 cuniculisepticus (Bacillus), 547, 652 cuniculicida (Bacillus) , 547, 552 cuniculicida (Bacterium) , 547, 681 cuniculicida (Pasteurella), 547 cuniculicida havaniensis (Bacillus), 612 cuniculicida havaniensis (Bacterium), 612 cuniculicida immobilis (Bacillus), 653 cuniculicida immobilis (Bacteriiim) , 653 cuniculicida mobilis (Bacillus) , 552 cuniculicida mobilis (Bacterium) , 552 cuniculicida thermophilus (Bacillus) , 681 cuniculicida thermophilus (Bacterium), 681 cuniculicida var. immobile (Bacterium), 653 cuniculorum (Micrococcus), 271 cunieuli (Leptotrichia) , 366 cupularis (Bacillus), 612 cupularis (Micrococcus) , 259 cupuliformans (Nanus), 1207 cupulijormis (Micrococcus) , 259 cursor (Bacillus), 716 curtissi (Micrococcus), 259 curvum (Bacterium), 188 curvum (Rhizobium) , 224 cuticularis (Bacillus), 677, 755 cuticularis (Bacterium), 677 cuticularis albus (Bacillus), 755 cutirubra (Pseudomonas), 110, 442 cutirubra (Serratia), 110 cutirubrum (Bacterium), 110 1422 INDEX OF NAMES OF GENERA AND SPECIES cuti-dare cutirubrum {Flavobacterium) {Halobac- terium), 110 cutis (Bacillus), 403 cutis {Corynebacterium) , 403 cutis commune (Bacterium), 403 cutis communis {Bacillus), 403 cutis communis (Micrococcus), 251, 259 cutis communis (Staphylococcus), 259 cyaneofluorescens (Bacillus), 145 cyaneofluorescens (Pseudomonas), 145, 146 cyaneo-fuscus (Bacillus), 233 cyaneophosphorescens (Achromobacter) 634 cyaneo-phosphorescens (Bacillus), 634 cyaneo-phosphorescens (Vibrio), 634 cyaneum (Bacteridium), 259 cyaneum (Photobacterium) , 634 cyaneus (Actinococcus) , 923 cyaneus (Bacterium), 695 cyanexis (Micrococcus) , 259, 272 cyaneus (Nigrococcus) , 259 cyaneus (Proactinomyces) , 923 cyaneus -antibioticus (Proactinomyces) , 923 cyanofuscus (Bacterium) , 233 cyanogenes (Pseudomonas), 92, 96 cyanogenes (Vibrio), 92 cyanogenum (Bacterium), 700 cyanogenus (Bacillus), 92, 96 cyanogenus (Micrococcus) , 259, 695 cyanoides (Agarbacterium), 630 cyanophos (Micrococcus) , 636 cyano -phosphor escens (Photobacterium) , 634 cycloclastes (Archromobacter), 420 cycloclastes (Bacterium), 420 Cyclops (Micrococcus), 259 cyclosites (Vibrio), 200 cygneus (Bacillus), 642 cygni (Bacterium), 642 cylindracea (Nocardia), 919 cylindracea (Oospora), 919 cylindraceus (Actinomyces), 919 cylindraceus (Discomyces) , 919 cylindricus (Bacillus), 731 cylindricus (Chondromyces), 1038 Cylindrogloea, 873 cylindroides (Bacterium) , 577 cylindroides (Ristella), 577 cylindrosporum (Clostridium), 789 cylindrosporus (Bacillus), 716, 718 cyprinicida (Bacterium), 642 cyprinicida (Klebsiella) , 642 cijpripedii (Bacillus), 470 cypripedii (Erivinia), 470 cystiformis (Bacillus), 653, 741 cystinovorum (Achromobacter) , 416 cystiopoeus (Micrococcus) , 259 cystitidis (Bacillus), 653 cystitidis (Streptococcus) , 339 Cystobacter, 1025, 1034 Cystodesmia, 14, 1036 Cystoecemia, 14, 1021 cytaseum (Bacterium) , 741 cytaseus (Bacillus), 741 cytaseus var. zonalis (Bacillus), 741 cytolytica (Erwinia), 473 Cytophaga, 35, 583, 1005, 1009, 1010, 1012 cytophaga (Spirochaeta) , 1049 cytophagus (Micrococcus), 259 cytophagus (Mycococcus), 1013 czaplewskii (Bacterium), 590 dacryoideus (Bacillus), 653 Dactylocoena, 14, 1044 dacunhae (Achromobacter), 105 dacunhae (Pseudomonas), 105 dadhi (Streptothrix) , 364 dahliae (Bacillus), 470 dahliae (Erwinia), 470 dahliae (Marmor), 1179 damnosus (Pediococcus) , 250 dainnosus (Streptococcus), 250 damnosus var. mucosus (Streptococcus) , 250 danicus (Bacillus), 714 danteci (Bacillus), 742 dantecii (Micrococcus), 259 danubica (Microspira) , 196 danubicum (Spirillum), 196 danubicus (Vibrio), 196 danysii (Salmonella), 517 dar-es-salaam (Salmonella) , 519 1423 dare deni INDEX OF NAMES OF GENERA AND SPECIES daressalaamensis (Salmonella) , 519 dassonvillei (Actinomyces), 915 dassonvillei (Discomyces) , 916 dassonvillei (Nocardia), 916 dassonvillei (Streptothrix) , 934 daucarum (Bacillus), 742 daxensis (Spirochaeta), 1053 daytona (Salmonella) , 513 de baryanus (Bacillus), 813 debile (Bacterium), 677 debilis (Streptococcus) , 339 debilitans (Legio), 1257 decalvens (Bacterium), 259 decalvens (Micrococcus) , 259 decidiosus (Bacillus), 441 decidiosus (Bacterium), 441 deciduosa (Cellulomonas) , 621 deciduosum (Flavobacterium) , 441 deciduosus (Bacillus) , 621 decipiens (Micrococcus), 259 decipiens (Pacinia), 696 decolor (Bacillus) , 653 decolor (Micrococcus) (Streptococcus 259 decolorans (Aerobacter) , 456 decolorans (Citrobacter) , 448 decolorans major (Bacillus) , 653 decolorans minor (Bacillus), 653 decussata (Nocardia), 975 decussata (Oospora), 975 decussatus (Actinomyces) , 975 decussatus (Discomyces) , 975 defessus (Bacillus) , 653 deformans (Micrococcus) , 259 deformans (Phagus), 1135 degenerans (Bacillus) , 637 degenerans (Microspira) , 637 degenerans (Photobacterium) , 637 dehydrogenans (Flavobacterium), 613 dehydrogenans (Micrococcus) , 613 delabens (Bacillus), 147 delabens (Bacterium) , 698 delabens (Pseudomonas) , 147, 697 delacourianus (Micrococcus) , 259 delbruckii (Bacillus), 355 delbruckii (Lactobacterium) , 355 delbrueckii (Lactobacillus) , 355, 695, delbruecki (Plocamobacterium) , 695 delbruecki (Ulvina), 695 delendae-muscae (Bacterium) , 677 delesseriae (Bacterium), 625 delesseriae (Flavobacterium) , 625 delgadense (Photobacterium), 637 delgadensis (Microspira) , 637 delicatulum (Achromobacter), 419 delicatulus (Bacillus), 419 delicatulus (Bacterium), 419 delicatum (Corynebacterium) , 403 deliense (Bacterium) , 677, 759 delmarvae (Achromobacter), 422 delphinii (Annulus), 1216 delphinii (Bacillus) , 115 delphinii (Bacterium) , 115 delphinii (Pectobacterium) , 696 delphinii (Phytomonas) , 115 delphinii (Pseudomonas), 115 de/^a (Bacillus), 653 de?to (Bacterium) , 653 demmei (Bacillus), 742 f), demmei (Micrococcus) , 270 dendriticum (Achromobacter), 423 dendriticus (Bacillus) , 423 dendriticus (Bacterium) , 423 dendrobii (Bacterium) , 613, 640 dendroides (Bacillus), 718, 742 dendroporthos (Micrococcus), 259 denekei (Vibrio), 196 denekii (Pacinia), 196 denieri (Alcaligenes) , 416 denitrificans (Bacillus), 423, 441, 442, 653, 754 denitrificans I (Bacillus) , 440 denitrificans II (Bacillus), 426 denitrificans (Bacterium) , 440, 688 denitrificans I (Bacterium) , 440 denitrificans (Chromobacterium) , 441 denitrificans (Flavobacterium) , 440, 688 denitrificans (Micrococcus), 260 denitrificans (Pseudomonas), 98 denitrificans (Sulfomonas) , 80 denitrificans (Thiobacillus), 80 denitrificans (Vibrio), 426 denitrificans agitis (Bacillus), 422 762 denitrificans agilis (Bacterium) , 423 1424 INDEX OF NAMES OF GENERA AND SPECIES deni-dick denitrificans fluorescens {Bacillus), 98 Denitrohacterium, 8 denitrofluorescens (Bacillus), 653 Denitromonas , 8, 83 dentale (Leptospira), 1078 dentalis viridans (Bacillus) , 653 dentahis (Bacillus), 742 denticola (Spirochaeta) , 1075 denticola (Spirochaete) , 1075 denticola (Treponema) , 1075 dentinum (Spirochaeta) , 1075 dentium (Fusiformis) , 581 dentium (Leptospira), 1079 dentium (Micrococcus) , 339 dentium (Spirillum), 1075 dentium (Spirochaeta), 1065, 1070, 1074, 1075 dentium (Spirochaete), 1075 dentium (Spironema) , 1075 dentium (Streptococcus) , 339 dentium (Treponema) , 1074 dentium-steogyratum (Treponema), 1075 deprimata (Cytophaga), 1013 derby (Salmonella) , 505 derbyensis (Salmonella) , 505 dermacentrophila (Rickettsia), 1096 dermacentroxenus (Rickettsia), 1087 dermatogenes (Micrococcus), 260 dermatogenes (Pseudomonas), 93 dermatonomus (Actinomyces), 916 dermoides (Bacillus), 653 dermophilum (Corynebacterium) , 403 dermophilus (Bacillus), 403 derossii (Bacillus), 803 derossii (Clostridium) , 803 desaiana (Phytornonas), 121 desaiana (Pseudomonas), 121, 125 desidens (Micrococcus), 339 desidens (Streptococcus), 339 desidiosa (Cellulomonas), 621 desidiosum (Flavobacterium) , 441 desidiosus (Bacterium), 441 desiduosis (Bacillus), 441, 621 desmodilli (Pasteurella) , 553 desmolyticum (Achromobacter) , 104 desmolyticum (Pseudomonas), 104 destillationis (Bacterium), 576 destillationis (Ristella), 576 destructans (Bacterium), 470 destructans (Phytornonas), 470 destructans (Pseudomonas) , 470 destruens (Bacillus), 742 Desulfovibrio, 29, 30, 35, 82, 207, 209 desulfuricans (Bacillus), 207 desulfuricans (Desulfovibrio), 207, 208, 209 desulfuricans (Microspira), 207 desulfuricans (Spirillum), 207 desulfuricans (Sporovibrio) , 207, 208 desulfuricans (Vibrio), 207, 208, 853 Detoniella, 983 detrudens (Bacillus), 742 devorans (Bacillus), 430 devorans (Bacterium), 430 devorans (Flavobacterium), 430 devorans (Sarcina), 291 devorans (Vibrio), 204 dextranicum (Leuconostoc), 347 dextranicum (Streptobacterium) , 701 dextranicus (Lactococcus) , 347 dextrolacticus (Bacillus), 712 diacetilactis (Streptococcus), 339 diacetyl aromaticus (Streptococcus) , 339 Dialister, 21, 27, 32, 33, 577, 594 Dialisterea, 20 dianthi (Bacillus), 640 dianthi (Bacterium) , 640 dianthi (Pseudomonas) , 640 diaphanus (Bacillus), 653 diaphthirus (Bacillus), 737, 793 diaporica (Rickettsia), 1092 diastasius (Thermobacillus) , 731 diastaticus (Actinomyces), 939 diastaticus (Bacillus) , 742 diastaticus (Streptomyces), 939 diastatochrojuogenes (Actinomyces) , 941 diastatochromogenes (Streptomyces) , 941 diatrypeticum (Bacterium), 677 diatrypeticus casei (Bacillus), 677 dichotoma (Cladothrix), 934, 975, 982 dichotoma (Nocardia), 975 dichotomus (Sphaerotilus), 982 dicksonii (Actinom.yces) , 969 1425 dicr-diss INDEX OF NAMES OF GENERA AND SPECIES Dicrohactnim, 13, 14, 479 Dicrospira, 12, 13, 192 Dicrospirilluni, 12, 13, 28, 212 didelphis (Spirochaeta) , 1066 Didymohelix, 23, 26, 29, 831 dieffenhachiae {Bacterium) , 157 dieffenhachiae (Phytomonas), 157 dieffenhachiae (Xanthomonas), 157 difficile (Clostridium), 773 difficilis (Bacillus), 773 diffluens (Bacillus), 490, 653 diffluens (Cytophaga), 1015 diffluens (Micrococcus), 260 diffluens (Proteus), 490 diffragens (Bacillus), 813 diffusum (Flavobacterium), 429 diffusus (Bacillus), 429 diffusus (Bacterium) , 429 digestans (Bacillus), 730 digitatus (Bacillus), 653 digitatus (Myxococcus), 1045 dihydroxyacetonicum (Bacterium), 189 dilaboides (Bacillus), 720 dilatator (Caryococcus), 1121 dimorpha (Mycoplana), 191 dimorpha (Urosarcin.a), 294 dimorphohutyricus (Bacillus), 814 dimorphus (Bacillus), 352 dimorphus (Bacteroides), 352 dimorphus (Micrococcus), 260 dimorphus var. longa (Bacillus), 352 diphtheria vitulorum (Oospora), 5ilS diphtheriae (Bacillus), 383 diphtheriae (Bacterium), 383, 401 diphtheriae (Corynebacterium), 383, 384, 385,406,752, 1131, 1143, 1144 diphtheriae (Phagus), 1143 diphtheriae avium (Bacillus), 400 diphtheriae avium (Bacterium) , 400 diphtheriae avium (Mycobacterium) , 915 diphtheriae columbarum (Bacillus), 401 diphtheriae columharum (Bacterium) , 401 diphtheriae cuniculi (Bacillus), 402 diphtheriae cuniculi (Bacterium) , 402 diphtheriae ulcerogenes cutaneum (Cory- nebacterium), 406 diphtheria vitulorum (Bacillus), 401, 578 diphthericum (Microsporon) , 383 diphthericus (Micrococcus), 260 diphtheritic us (Streptococcus) , 260, 337 diphtheroides (Bacillus), 401 diphtheroides (Coccobacillus), 402 diphtheroides (Corynebacterium), 388, 403 diphtheroides (Kokkobacillus) , 402 diphtheroides brevis (Bacillus), 402 diphtheroides citreus (Bacillus), 406 diphtheroides gallinarum (Bacillus), 403 diphtheroides liquefaciens (Bacillus), 404, 405 Diplectridium, 1 , 763 Diplobacillus , 590 Diplococcus, 13, 17, 20, 31, 33, 42, 305 Diplostreptococcus , 312 disciformans (Bacillus), 368, 742 disciformans (Bacterium), 742 disciformans (Eubacterium) , 368 disciformis (Angiococcus), 1047 disciformis (Bacillus), 742 disciformis (Myxococcus), 1047 discofoliatus (Actinomyces), 927 discoides (Neisseria), 299 Discomyces, 925 discophora (Chlamydothrix) , 985 discophora (Leptothrix), 985 discophora (Megalothrix) , 985 dispar (Actinomyces), 918 rf/spor (Bacillus), 540, 542, 543 dispar (Bacterium), 542 dispar (Discomyces) , 918 dispar (Eberthella) , 42, 542 dispar (Eperythrozoon), 1113 dispar (Microsporon), 918 dispar (Proshigella) , 542 dispar (Shigella), 542 dispar (Sporotrichum) , 918 disparis (Streptococcus), 339 Dispora, 349 disporum (Clostridium) , 820 dissimilis (Bacillus), 653 dissimilis (Micrococcus) , 260 dissolveiis (Aerobacter) , 472 dissolvens (Aplanobacter) , 472 dissolvens (Bacterium) , 472 dissolvens (Clostridium), 43, 809, 810 1426 INDEX OF NAMES OF GENERA AND SPECIES diss-dupl dissolvens (Erwinia), 464, 472 dissolvens {Phytomonas), 472 dissolvens {Pseudomonas), 472 Distasoa, 20, 21, 23, 27 distasonis (Bacteroides), 570 distasojiis (Ristella), 570 distendens (Streptococcus), 347 distortus (Bacillus), 742 distortus (Tyrothrix), 742 diversum (Acetobacter) , 692 diversum (Aerobacter) , 456 diversum (Citrobacter) , 448 djokjakartensis (Micrococcus), 260 dmitrovi (Spirochaeta), 1078 dobelli (Bacillus), 742 dobelli (Bacillus) (Flexilis), 742 dodecahedron (IMarmor), 1169 doederlein (Lactobacillus) , 362 doederleinii (Acidobacterium), 362 dornesticus (Bacillus) , 653 domesticus (Bacterium), 653 donnae (Actinomyces), 916 Donovania, 559 dorz (Actinomyces), 916 (fori (Discomyces), 916 do/-i (Nocardia), 916 don (Oospora), 916 don (Rhinocladium) , 916 don (Sporotrichum) , 916 doriae (Oospora), 934, 968 dormitator (Bacillus), 437 dorrnitator (Bacterium), 437 dormitator (Flavobacterium), 437 douglasi (Bacillus), 544 douglasi (Shigella), 544 Douglasillus, 11, 763 dowdesivelli (Urococciis), 282 doyeni (Bacillus), 651 doyeni (Micrococcus) , 260 drennani (Vibrio), 204 Drepanospira, 1122 drimophylus (Micrococcus), 260 droebachense (Bacterium), 625 droebachense (Flavobacterium) , 625 droebachense (Pseudomonas), 625 droserae (Bacillus), Q53 droserae (Bacterium), 653 drosophilae (Treponema), 1075 dschunkowskii (Grahamella), 1110 dschunkowski (Grohamia), 1110 dubia (Eberthella) , 533 dubitata (Palmula), 812 dubitatus (Acuformis), 812 dubium (Bacterium), 703 dubium (Marmor), 1172, 1214 dubium (Rhizobium), 225 dubius (Annulus), 1155, 1214 dubius (Bacillus), 533 dubius (Bacterium), 533 dubius (Phagus), 1137 dubius pneumoniae (Bacillus), 703 dubius var. annulus (Annulus) , 1215 dubius var. flavus (Annulus) , 1216 dubius var. obscurus (Annulus) , 1216 dubius var. vulgaris (Annulus), 1215 dublin (Salmonella) , 517 dublin var. accra (Salmonella) , 517 dublin var. ^oeZn. (Salmonella) , 517 duclauxii (Bacillus), 654, 742 duclauxii (Urobacillus), 654, 688, 729, 742 ducreyi (Coccobacillus), 587 ducreyi (Hemophilus), 587 dudtschenkoi (Grahamella), 1110 duesseldorf (Salmonella) , 514 dulcito-fermentans (Bacillus), 772, 824 dunbari (Microspira) , 203 dunbari (Photobacterium) , 203 dunbari (Photospirillum) , 203 dunbari (Vibrio), 702 duodenale (Bacterium), 447 duodenale (Encapsulata) (Bacillus), 447 duplex (Bacillus), 590 duplex (Bacterium) , 590, 834 duplex (Ferribacterium) , 834 duplex (Hemophilus), 591 duplex (Moraxella) , 592 duplex (Pseudomonas) , 147 duplex (Siderobacter) , 834 duplex (Siderocystis) , 835 duplex (Sideroderma) , 834 duplex josephi (Bacillus), 592 duplex josephi (Moraxella), 592 duplex liquefaciens (Bacillus), 591 duplex liquefaciens (Moraxella), 591 1427 dupl-elus INDEX OF NAMES OF GENERA AND SPECIES duplex non-liquejaciens {Bacillus), 592 duplex non-liquefaciens (Bacterimn), 592 duplex non liquefaciens {Moraxella), 592 duplex var. liquefaciens (Moraxella), 591 duplex var. non liquefaciens (Moraxella), 592 duplicatus (Bacillus), QdS duplicatus (Bacterium), 693 durabilis (Phagus), 1142 durans (Streptococcus), 327 durhan (Salmonella), 519 duttoni (Cacospira), 1060 duttoni (Spirillum), 1060 duttoni (Spirochaeta), 1060 duttoni (Spirochaeta) (Microspironema), 1060 duttoni (Spironema), 1060 duttoni (Spiroschaudinnia), 1060 duttoni (Treponejna), 1060 duttonii (Borrelia), 1060, 1061, 1064, 1066 dysenteriae (Bacillus), 536, 537, 538, 539, 540 dysenteriae (Bacterium), 536, 537, 689 dysenteriae (Eberthella) , 536 dysenteriae (Phagus), 1132 dysenteriae (Shigella), 535, 536, 537, 542, 1131, 1132, 1133, 1134, 1135 dysenteriae Boyd I to /// (Bacillus), 538 dysenteriae Flexner I to VI (Bacillus) , 538 dysenteriae Flexner VTI and YIII (Bacil- lus), 538 dysenteriae liquefaciens (Bacillus), 543 dysenteriae liquefaciens (Bacterium), 543 dysenteriae Schmitz (Bacillus), 536 dysenteriae vitulorum (Bacillus), 689 dysenteriae vitulorum (Bacterium) , 689 dysentericus (Bacillus), 536, 543 Dysenteroides, 10 dysgalactiae (Streptococcus), 319 dysodes (Bacillus) , 742 eastbourne (Salmonella) , 519 eatonii (Micrococcus), 260 Eberthella, 10, 21, 26, 37, 42, 494, 516, 533 Eberthella sp. (Sendai type), 51S Eberthus, 10, 516 eburneus (Micrococcus), 260 echinata (Leptothrix), 985 eczemae (Micrococcus) , 278 eczemicus (Bacillus), &5A cdematis (Clostridium), 775 edgeworthiae (Bacillus), 478 edgeworthiae (Erwinia), 478 edigtoni (Neisseria), 301 efficiens (Borrelina), 1226 efficiens (Marmor), 1189, 1191 effrenus (Phagus), 1143 effusa (Cellulomonas) , 91 effusa (Pseudomonas), 91 effusavar.nonliquefaciens (Pseudomonas) , 92 cffusus (Bacillus), 91, 718 egens (Bacillus), 790 cgfens (Clostridium), 790, 826 egens (Stoddardillus) , 790 egregius (Bacillus), 654 egypticum (Treponema), 1065 Ehrenbergia, 37, 1052 ehrenbergii (Bacillus), 597 ehrenbergii (Bacterium) , 597 Ehrlichia, 1096 ehrlichii (Grahamella) , 1110 Eisenbergia, 12, 13, 486, 705 eisenbergii (Pseudomonas), 97, 98, 698 elaphorum (Bacterium) , 145 elastica (Actinomyces), 969 elegans (Bacillus), 742, 814 elegans (Flexibacter) , 38 elegans (Holospora), 1122 elegans (Thiodictyon), 845, 849 elipsoideus (Bacillus), 654 ellenbachensis (Bacillus), 715 ellenbachensis alpha (Bacilhis) , 715 ellenbachi (Bacillus), 717 ellingeri (Coccobacillus), 452 ellingeri (Escherichia), 452 ellingtonii (Bacillus), 654 ellipsoidea (Pseudomonas), 147 ellipsospora (Cytophaga), 1050 ellipsospora (Sporocytophaga), 1050 elongata (Pseudomonas), 698 elongata (Thiospira), 702 eZwsa (Spirochaeta), 1079 1428 INDEX OF NAMES OP GENERA AND SPECIES elus-epid elusum (Treponema) , 1079 eminans (Bacillus), 654 emphysematis maligni (Bacillus), 791 emphysemaiis vaginae (Bacillus), 790, 826 emphysematosus (Bacillus), 789, 790 emphysematosus (Bacterium), 790 emulsinus (Bacillus), 654 emulsionis (Bacillus), 742 enalia (Pseudomonas) , 698 Encapsulatus, 10, 17, 18, 457 encephaloides (Bacillus), 742 enchelys (Bacillus), 677 enchelys (Bacterium) , 677 endiviae (Phytomonas) , 133 endiviae (Pseudomonas) , 133 Endobacterium, 705 endocarditicus (Streptococcus), 339 endocarditidis (Bacillus), 654 endocarditidis (Bacterium), 677 endocarditidis capsulatus (Bacillus), 677 endocarditidis griseus (Bacillus), 654 endocarditidis griseus (Bacterium), 654 endocarditis (Cillohacterium) , 369 endocarditis (Micrococcus), 274 endocarditis griseus (Bacillus), 401 endocarditis rugatus (Micrococcus), 274 endometritidis (Bacillus), 677 endometritidis (Bacterium), 677 endometritis (Plocamobacterium) , 677 endometritis canis (Bacterium), 677 endoparagogicum (Spirillum), 217 Endosporus, 33, 34, 763 englemanni (Bacillus), 654 enterica (Eberthella) , 533 enterica (Escherichia), 450 entericus (Bacillus), 450, 533 entericus (Enteroides) , 450, 534 entericus (Proteus), 489 enteritidis (Actinomyces), 920 enteritidis (Bacillus), 505, 516, 517, 782, 818, 920 enteritidis (Bacterium), 516 enteritidis (Discomyces), 920 enteritidis (Klebsiella), 510 enteritidis (Nocardia), 919 enteritidis (Oospora), 920 enteritidis (Phagus), 1136 enteritidis (Salmonella), 493, 497, 516, 517, 523, 531, 1130, 1136, 1137 enteritidis (Streptococcus), 339 enteritidis (Sf.revtothrix), 919, 976 enteritidis breslau (Bacterium), 502 enteritidis breslaviense (Bacillus), 502 enteritidis sporogenes (Bacillus), 782, 818 enteritidis sporogenes (Clostridium), 818 enteritidis-yellow (Salmonella), 531 enteritidis var. c/iaco (Salmonella), 517 enteritidis var. danysz (Salmonella), 517 enteritidis var. dublin (Salmonella), 517 enteritidis var. esse?i (Salmonella), 517 enteritidis var. jena (Salmonella), 517 enteritidis var. moscow (Salmonella) , 518 enteritidis var. miilheim (Salmonella) , 517 enteritidis var. rostock (Salmonella) , 518 enteritidis B, Typ. equinus (Bacillus), 506 enteritidis B,Typ. murium (Bacillus), 502 enteritidis C, Typ. oz;is (Bacillus), 506 enteritidis var. F (Salmonella), 531 enteritis (Streptococcus) , 339 enteritis var. libmanii (Streptococcus), 339 Enterobacter , 31, 32, 37 Enter obacterium, 37 Enterococcus , 326, 336 cnterococcus (Diplococcus) , 325 enterocoliticum (Bacterium), 677 Enteroides, 10 enteroideus (Micrococcus) , 695 enteromyces (Bacillus), &54: enterothrix (Bacillus), 742 Entomospira, 12, 13, 28, 1058 entomotoxicon (Bacillus), 654 enzymicum (Corynebacterium), 386, 407 enzymicus (Bacillus) , 386 enzymothermophilus (Lactobacillus), 363 cos (Mycobacterium) , 905 Eperythrozoon, 1100, 1111, 1113 Eperythrozoon spp., 1113 ephemerocyanea (Pseudomonas), 147 ephestiae (Micrococcus), 260 ephestiae No. 1 and A^o. ;? (Bacterium), 759 epidemicus (Streptococcus), 315 epidermidis (Albococcus) , 243 1429 epid-erys INDEX OF NAMES OF GENERA AND SPECIES epidermidis (Bacillus) , 742 epidermidis {Corxjnebacterium) , 403, 406 epidermidis (Leptothrix), 691, 742 epidermidis (Micrococcus), 243, 252, 254, 255, 256, 259, 264, 265, 270, 271, 272 epidermidis {Plocamohacterium) , 691 epidermidis (Staphylococcus), 243 epidermidis var. A (Albococcus) , 243 epidermidis albus (Micrococcus) , 243, 268 epidermidis albus (Staphylococcus), 243 epidermis (Micrococcus) , 278 Epidermophyton sp., 921 epimetheus (Micrococcus), 260 epiphitica (Chlamydothrix) , 986 epiphytica (Leptothrix), 985 epiphytica (Lyngbya), 986 epiphytica (Streptothrix) , 885 epiphytus (Bacillus) , 743 eppingeri (Actinomyces) , 896 eppingerii (Streptotrix) , 896 epsilon (Bacillus), 654 epsilon (Bacterium), 654 epsteinii (Achromobacter), 424 equae (Tortor) , 1278 equarius (Streptococcus) , 339 equatilis communis (Bacillus). 102 equestris (Malleomyces), 554 equi (Actinomyces) , 92{) equi (Bacillus), 654 equi (Bacterium), 541 ejuz (Bollingera) , 253 egwi (Botryomyces), 253 equi (Corynebacterium), 391 egwi (Corynethrix), 406 e^Mt (Discomyces) , 252, 920 egw (Mycobacterium) , 391 egwi (Nocardia), 920 egwi (5amna), 290, 291, 292, 294 egwz (Shigella), 541 egwi (Spirillum), 1066 e^Mi (Spirochaeta) , 1066 eguz (Spironema) , 1066 egwi (Spiroschaudinnia) , 1066 equi (Streptococcus), 317, 31S egu'j (Treponema), 1066 eg-m intestinalis (Bacillus) , 654 eg«i iyitestinalis (Bacterium), 654 equidistans (Bacillus) , 1098 equina (Spirochaeta), 1066 equinus (Erro), 1253 equinus (Hostis), 1240 equinus (Streptococcus), 323, 339 equirulis (Bacillus), 541 equirulis (Shigella), 540 equiseptica (Pasteurella), 553 equisepticus (Bacillus), 553 equisimilis (Streptococcus), 318, 319 equorum (Tortor), 1277 equorum (Trifur), 1282 equuli (Bacillus), 541 erebea (Legio), 1259 erectum (Podangium), 1008, 1034 erectus (Chondromyces), 1034 erectus (Cystobacter), 1034 eriobotryae (Bacterium), 144 eriobotryae (Phytomonas) , 144 eriobotryae (Pseudomonas), 144 erivanense (Bacterium) , All erivanensis (Bacillus), 471 erivanensis (Erwinia), 471 Ermengemillus, 11, 763 erodens (Bacillus), 743 erodens (Foliopellis), 1171 erodens (Marmor), 1171 erodens var. severum (Murmur), 1171 erodens var. vulgare (Marmor), 1171 erodii (Bacterium) , 121 erodii (Phytomonas), 121 erodii (Pseudomonas), 121 Erro, 1248 erubescens (Bacillus), 654 Erwinia, 31, 42, 443, 463, 476 erysipelatis (Micrococcus), 315 erysipelatos (Staphylococcus), 315 erysipelatos (Streptococcus), 315, 1139 erysipelatos -suis (Bacillus), 410 erysipelatos-suis (Erysipelothrix), 410 erysipelatos suum (Bacterium), 410 erysipelatosus (Streptococcus), 315 erysipeloides (Babesia), 411 erysipeloides (Streptothrix) , 411 erysipeloidis (Actinomyces), 411 erysipeloidis (Bacterium) , 411 1430 INDEX OF NAMES OF GENERA AND SPECIES erys-€xan erysipeloidis (Erysipelothrix), 411 erysipeloidis (Oospora) , 411 Erysipelothrix, 18, 21, 22, 27, 28, 35, 37, 38, 409, 410 erythematis {Bacillus) , 742 erythematis {Bacterium) , 742 erythematis maligni {Bacillus) , 742 erythra {Pseudomonas), 147 erythraeus {Bacillus), 641 erythrea {Nocardia), 920 erythrea {Stre-ptothrix) , 920 erythreus {Actinomyces) , 938 erythreus (Streptomyces), £38 Erythrobacillus, 10, 479 Erythrobacterium, 32 erythrochromogenes {Actijiomyces) , 944 erythrochromogenes {Streptomyces), 944 Erythroconis, 844 erythrogenes {Bacillus), 600 erythrogenes (Bacterium), 600, 601, 602 erythrogenes {Corynebacterium) , 600 erythrogenes {Erythrobacillus) , 600 erythrogenes rugatus {Bacillus), 654 erythrogloeum (Bacterium), 637 erythromyxa {Bacillus) , 677 erythromyxa {Bacterium), 677, 687 erythromyxa {Micrococcus) , 291 erythromyxa {Micrococcus) {Staphylo- coccus), 392, 677 erythromyxa {Rhodococcus) , 8, 677 erythromyxa {Sarcina), 291 erythropolis {Actinomyces) , 898 erythropolis {Mycobacterium), 898 erythropolis (Nocardia), 898 erythropolis {Proactinomyces) , 898 erythrospora {Pseudomonas) , 147 erythrospores {Bacillus) {Streptobacter) , 654 erythrosporus {Bacillus), 147, 654 erythrosporus {Bacterium) , 654 erzinjan {Salmonella) , 507 Escherichia, 3, 10, 21, 26, 31, 37, 443, 444, 448, 450, 458, 492, 694 escherichii {Bacillus), 445 essen 773 {Salmonella) , 505 esseyana {Serratia), 484 esterificans {Bacillus), 43, 743 ester ijicans {Micrococcus), 260 esterificans {Plectridium) , 743 esterificans flaorescens {Bacillus), 654 esterificans stralauense {Bacterium), 654 esteroaromaticum {Bacterium) , 436 esteroaromaticum (Flavobacterium), 436 e|/ .Ci sV ^ r^s ^;>>j^ '

n anaerobicum (Coccobacterium) , 511 mucosum capsulatum (Bacterium) , 459 mucosus (Actinomyces), 916 7nucosus (Bacillus), 749, 788 7nucosus (Capsula7-is), oil 771UCOSUS (Cellulobacillus) , 762 mucosus (Diplococcus), 301, 308 77UICOSUS (Endosporus), 788 7nucosus (Mycococcus), 891 mucosus (Pneumococcus) , 301, 308 /nucosus (Streptococcus), 301, 308, 1139 771UCOSUS anaerobius (Bacillus) , 511 mucosus capsulatus (Bacillus) , 458, 663 771UCOSUS capsulatus (Streptococcus) , 308 /71UC0SUS ozaenae (Bacillus), 459 7nucosus ozaena (Bacte7'ium) , 459 77iucronatus (Bacillus) , 749 muelleri (Caryophanon), 1004 muelleri (Simonsiella) , 1004 i7menchen (Sal77ionella), 513 muenster (Sahjionella) , 523 77iulieris (Vibrio), 205 mulleri (Achro7natiu77i) , 1000 miilleri (Drepanospira), 1122 1469 mull-musc INDEX OF NAMES OF GENERA AND SPECIES mulleri (Monas), 1000 mulleri {Thiovulum) , 1000 Multifermentans , 11, 763 multif ermentans (Clostridium) , 772 multifermentans tenalbus (Bacillus), 772 multiforme (Cillobacterium) , 369, 790, 815 multiforme psittacosis (Microbacteriuin), 1116 multiformiis (Heterocysiia) , 13 multiformis (Bacillus), 369, 663, 790, 815, 826 multiformis (Bacteroidea) , 22 multiformis (Bacteroides) , 790, 815 multiformis (Haverhillia) , 588, 972 multiformis trichorrhexidis (Bacillus), 688 multipediculum (Bacterium) , 682 multipediculus (Bacillus), 682 multipediculus flavus (Bacillus), 685, 749 multistriata (Pseudomonas), 101, 699 multistriatum (Achromobacter) , 101 multistriatus (Bacillus) , 101 multistriatus (Bacterium), 699 inultivolaligenum (Lactobacterium) , 363 multocida (Pasteurella) , 546, 549 multocidum (Bacterium), 546, 547 muralis (Bacillus) , 749 muricida (Pasteurella), 548 muricola (Rickettsia), 1086 murina (Rickettsia), 1086 murinus (Bacillus), 411, 663 muripestifer (Bacillus), 682 muripestifer (Bacterium) , 682 muris (Actinomyces), 588, 972, 1109 muris (Asterococcus) , 588 muris (Bacillus), 402 muris (Bacterium) , 401 muris (Bartonella), 1103 muris (Borrelia), 215 muris (Fusiformis), 583 muris (Grahamella) , 1110 muris (Haemobartonella), 1103, 1104, 1105, 1106, 1107, 1108 muris (Hemophilus), 589 muris (Legio), 1261 mwrn (Mycobacterium) , 890 muris (Nocardia) , 924 muris (Spirella), 215 muris (Spirochaeta) , 215 muris (Spironema), 215 muris (Treponema), 215 muris (Treponemella) , 215 muris musculi iberica (Grahamella) , 1110 muris musculi var. albinoi (Bartonella) , 1108 inuris musculi var. albinoi (Haemobar- tonella), 1108 muris ratti (Actinomyces), 588, 924, 972 muris ratti (Bartonella) , 1103 muris-ratti (Streptothrix) , 924, 972 muris var. galatziana (Spirochaeta) , 215 muris var. virginiana (Spirochaeta), 215 muris var. virginiana (Spironema), 215 muriseptica (Erysipelothrix), 411, 548 muriseptica (Pasteurella), 411, 548 muriseplicum (Bacterium), 411 murisepticum (Corynebacterium), 390 muriseplicum (Mycobacterium), 411 murisepticus (Bacillus) , 411, 548, 665 murisepticus (Streptococcus), 341 murisepticus pleomorphus (Bacillus) , 490, 665 murisepticus pleomorphus (Bacterium), 665 murium (Bacillus), 502 murium (Corynebacterium) , 390 murmanensis (Microspira) , 202 musae (Bacillus), 137 musae (Bacterium?) , 613 musae (Phytomonas?) , 613 musae (Pseudomonas), 613 musarum (Bacillus), 137 muscae (Staphylococcus), 282, 1142 muscoides (Bacillus), 815 muscoides (Cornilia) , 815 muscoides colorabilis (Bacillus), 815 muscoides non colorabilis (Bacillus), 815 muscorum (Chondromyces) , 1020 muscorum (Stelangium), 1020 musculi (Bacillus) , 714 musculi (Grahamella), 1110 musculi (Gyromorpha) , 1112 1470 INDEX OF NAMES OF GENERA AND SPECIES musc-natt musculorum {Actinomyces), 972 musculorum, suis {Actinomyces), 972 musculorum suis {Oospora), 972 mustelae {Bacillus) , 552 mustelae septicus {Bacillus), 445, 552 mustelaecida {Bacillus), 552 muslelaecida {Pasteurella) , 552 mutabile (Bacterium), 603 mutabile {Rhizobium), 224 mutahilis {Bacillus), 749 mutans {Streptococcus), 342 myceticus {Micrococcus), 268, 696 Mycobacterium, 7, 17, 18, 19, 23, 27, 28, 30, 35, 38, 392, 407, 863, 875, 876, 887, 891, 892, 903, 905 Mycobacterium spp., 882, 887 Mycococcus, 875, 891 Mycoderma, 9, 179 mycodermatus {Micrococcus), 268 mycodermatus {Tetracoccus) , 284 Mycogallionella, 986 mycogenes {Bacillus), 663 mycogenum {Bacterium), 663 mycoides (Asterococcus), 1259, 1291 mycoides (Bacillus), 612, 718, 719, 742 982, 1130, 1138 mycoides (Bacterium), 602 mycoides corallinus (Bacillus), 644, 902 mycoides peripneumoniae {Coccobacillis) , 1291 mycoides roseum {Bacterium), 602 mycoides roseum {Chromobacterium) , 602 mycoides roseus {Bacillus) , 602 mycoides-roseus {Erythrobacillus) , 602 mycoides var. ovoaethylicus {Bacillus), 720 Mycomonas, 8, 876 Myconostoc, 6 Mycoplana, 82, 191 Mycoplasma, 1291 Mycothrix, 983 myricae {Actinomyces), 972 mytili {Bacillus), 663 Myxobacillus, 705 Mrjxobacter, 1005, 1025 Myxobotyrs, 1036 myxococcoides {Cytophaga), 1049 myxococcoides (Sporocytophaga) , 1049 Myxococcus, 14, 17, 20, 24, 25, 312, 1007, 1009, 1040, 1041, 1044, 1047, 1049 myxodens {Bacillus), 749 myxogenes {Cellulobacillus) , 762 myxogenes {Clostridium), 820 myxogenes (Pseudomonas), 93 mj'xomae (Molitor), 1245 nacreaceum {Bacterium) , 682 nacreaceus {Bacillus), 682 nacreaceus {Micrococcus) , 268 nadsonii {Proteus), 491 naganoi {Sarcina), 293 naganophila {Spirochaeta) , 1068 nakatae {Phytomonas) , 164 nakatae (Xanthomonas), 164, 1136 nakatae type B {Bacterium) , 164 nana {Saprospira) , 1055 nanukayami {Spirochaeta), 1077 Nanus, 1206 nanus {Bacillus), 718, 815 naphthalinicus {Bacterium), 682 naphthalinicus liquefaciens {Bacillus) , 663 naphthalinicus non-liquefaciens {Bacil- Zms),663 napi (Savoia), 1221 napi {Vibrio), 205 napoli {Salmonella), 522 narashino {Salmonella) , 514 nasale {Spirillum), 203 nasale {Spirosoma) , 203 nasalis {Bacterium) , 459 nasalis {Micrococcus) , 342 nasalis {Planococcus) , 342 nasalis {Streptococcus), 342 7iasalis {Vibrio), 203 nasicola {Spirillum), 203 natans {Bacillus), 718 natans {Cladothrix) , 982 natans {Cryptococcus) , 258 natans (Sphaerotilis), 982 natans var. cladothrix {Sphaerotilus) , 982 nathansonii {Thiobacterium) , 81 naWo {Bacillus), 710 1471 Naum-nico index of names of genera and species N aumanniella , 834 navicula {Annjlohacter) , 111 navicula (Bacillus), 771 navicula {Bacterium), 771, 824 naviculum {Clostridium) , 771 naviformis (Bacillus), 576, 723 navijormis (Ristella), 576 n'dianka (Vibrio), 205 neapolitana (Escherichia), 447 neapolitamis (Bacillus), 447 neapolitanus (Bacterium) , 447 nebulosa (Pseudomonas), 101, 699 nebulosum (Achromobacter) , 101 nebulosus (Bacillus), 101, 580, 663, 749, 815 nebulosus (Bacterium), Q99 nebulosus (Cryptococcus), 258 nebulosus (Protococcus) , 258 nebulosus gazogenes (Bacillus), 749 necans (Bacillus), 663 necrodentalis (Bacillus), 361, 362 necrogenes (Bacillus), 579 necrogenes (Spherophorus) , 579 necrophora (Pasteurella) , 554 necrophora (Strcptolhrix) , 578, 977 necrophorum (Bacterium), 578 necrophorum (Cor^jnebacterium), 578 necrophorus (Actinomyces), 578 necrophorus (Bacillus), 578 necrophorus (Fusiformis), 578, 583 necrophorus (Spherophorus), 578, 580, 583, 928 necrosans (Clostridium), 820 necroseos (Bacillus), 578 necroseos (Streptococcus), 342 necroticans (Micrococcobacillus) , 690 necroticus (Bacillus), 579 necroticus (Spherophorus), 579 necroticus (Streptococcus), 342 nectarophila (Bacterium), 134 nectarophila (Phytomonas) , 134 nectarophila (Pseudomonas), 134 neddini (Actinomyces) , 916 negombensis (Bacillus), 544 negombensis (Shigella) , 544 neigeux (Bacille), 777 neisseri (Pacinia), 386 Neisseria, 19, 21, 26, 27, 29, 31, 33, 295 nelliae (Bacillus), 478 nelliae (Erwinia) , 478 nenckii (Achromobacter) , 624 nenckii (Bacterium) , 624 neocistes (Vibrio), 199 neoformans (Micrococcus) , 268 neotropicalis (Borrelia) , 1068 neotropicalis (Spirochaeta) , 1064 neotropicalis (Treponema), 1064 nephriticus (Bacillus) , 663 nephritidis (Bacterium), 553, 760, 761 nephritidis equi (Bacillus), 540 nephritidis instertitialis (Bacillus), 760 neptunium (Flavobacterium), 432 neritica (Pseudomonas), 699 nerviclarens (Marmor), 1198 neschezadimenki (Actinomyces), 928 neschezadimenki (Cohnistreptothrix) , 928 neumanni (Bacillus), 681, 749 iieurolyticus (Musculomyces), 1293 neurotomae (Bacillus), 663 neurotomae (Bacterium) , 663 neurotomae (Micrococcus), 268 neustonica (N aumanniella) , 834 iieuvillei (Micrococcus) , 268 neveuxi (Spirochaeta) , 1059 iieveuxi (Spironema) , 1059 neveuxi (Treponema) , 1059 neveuxii (Spiroschaudinnia), 1059 Xevskia, 7, 35, 829 A/ew brunswick (Salmonella), 525 newington (Salmonella), 524 neioport (Salmonella) , 573 newportensis (Salmonella) , 513 newport var. kottbus (Salmonella), 513 Newskia, 7, 12, 14 net« yor/j (Salmonella), 522 nexibilis (Bacillus), 148 nexibilis (Bacterium) , 699 nexibilis (Pseudomonas), 148, 699 nexifer (Micrococcus), 696 nicolaieri (Bacterium), 682 nicolaieri (Pacinia), 798 Nicollaierillus, 11, 763 nicollei (Actinomyces), 921 nicollei (Bartonella), 1108 1472 INDEX OF NAMES OF GENERA AND SPECIES mco-noca nicollei (Haemobartonella) , 1108 nicollei (Nocardia), 921 nicollei (Spirochaeta) , 1059 nicollei {Spironema) , 1059 nicollei {Treponema), 1059 nicomosaicum (Phytovirus) , 1164 nicotianae (Bacillus), 137 nicotianae (Erwinia), 137 nicotianae-tabaci {Phytomonas), 639 nicotianum (Bacterium), 682 nicotinobacter (Bacterium), 682 nictoinophagum (Bacterium) , 682 nictoinovorum (Bacterium) , 613 ni^e/- (Actinomyces) ,9(^9 niger (Bacillus) , 711 niger (Micrococcus), 247 nigra (Nocardia), 921 nigra (Streptothrix) , 921, 93-4, 969, nigrescens (Bacillus), 749 nigrescens (Micrococcus) , 268 nigrescens (Nigrococcus) , 268 nigrescens (Sorangium), 1024 nigricans (Actinomyces) , 972 nigricans (Bacillus), 749 nigrifaciens (Pseudomonas), 109 nigrificans (Actinomyces), 972 nigrificans (Bacillus), 711 nigrificans (Clostridium), 802 nigrificans (Oospora), 972 Nigrococcus, 10 nigrofaciens (Micrococcus) , 268 nigromaculans (Bacterium), 168 nigromaculans (Phytomonas), 168 nigromaculans (Xanthomonas), 168 nigrum (Catenabacterium) , 368 nigrum (Sorangium), 1024 nigrum (Spirillum) , 217 nijibetsui (Achromobacter) , 425, 692 nili (Erro), 1251 niloese (Salmonella), 525 nimipressuralis (Erwinia), 472 ninas kohl-yakomovi (Grahamella), 1110 niosii (Bacteroides) , 367 niosii (E ubacterium) , 367 nipponica (Rickettsia), 1091 nipponica (Rickettsoides) , 1091 nitens (Bacillus), 663 nitens (Bacterium), 682 nitidus (Arthromitus), 1003 nitidus (Bacillus), 749 nitidus (Micrococcus) , 269 nz (Bacillus), 653 oogenes hydrosulfureus s (Bacillus), 659 oogenes (Pseudomonas), 148 oonergasius (Bacillus) , 664 opaca (Nocardia), 897 opacum (Mycobacterium) , 897 opacum (Nitrobacter), 76 opacus (Bacillus), 750 opacus (Proactinomyces) , 897, 898 opacus (Streptococcus), 342 opalanitza (Leuconostoc) , 346 opalescens (Micrococcus) , 252, 270 Ophidomonas, 6, 850 Ophiocystia, 14, 1017 opossum (Bartonella), 1108 opossu7n (Haemobartonella), 1108 opportunus (Streptococcus), 342 orae (Annulus), 1155, 1212, 1213 orae (Bacillus), 750 oroe (Tractus), 1213 orangea (Sarcina), 293 orangica (Nocardia), 976 orangica (Slreptothrix) , 976 orangico-niger (Actinomyces), 972 orangicus (Actinomyces) , 972 orangium (Chromobacterium) , 694 oranienburg (Salmonella) , 510 oranienburgensis (Salmonella), 510 orbicularis (Micrococcus), 270 orbicularis Jlavus (Micrococcus), 270 orbiculata (Neisseria), 300 orbiculatus (Micrococcus), 270 orbiculus (Diplococcus), 300 orchiticum (Bacterium), 682 orchiticus (Bacillus), 683 orchitidis (Flavobacterium) , 556 Oregon (Salmonella) , 513 orenburgii (Bacillus), 362 orientate (Bacillus) , S&2 orientalis (Dennacentroxenus) , 1090 orientalis (Nanus), 1206 orientalis (Rickettsia), 1089, 1090 orientalis (Salmonella), 528 orientalis var. schuffneri (Rickettsia), 1090 orientuni (Propionibacterium), 379 orioAi (Salmonella), 524 orleanense (Bacterium), 187, 693 orleanensis (Ulvina), 693 ornithopi (Bacillus), 224 ornithosis (Miyagawanella), 1117, 1118, 1119 orpheus (Bacillus), 724 orthobutylicus (Bacillus), 111, 824 orthodonta (Spirochaeta), 1075 orthodontum (Treponeyna) , 1075 oryzae (Bacterium), 168 oryzae (Fractilinea), 1160 oryzae (Marmor), 1160 oryzae (Phytomonas), 168 oryzae (Pseudomonas), 168 oryzae (Xanthomonas), 168 Oscillatoria, 988, 991, 1007 Oscillospira, 1004 osio (Salmonella), 511 osteomyeliticus (Bacillus) , 664 osteomyelitus bubalorum (Bacillus), 778 osteophilum (Bacterium), 683 1476 INDEX OF NAMES OF GENERA AND SPECIES ostr-pall ostreae {Cristispira), 1056 ostreae (Spirillum), 217 ostreae (Spirochaeta), 1056 ostrei {Bacillus) , 532 ostrei (Salmonella) , 532 olitidis sporogenes putrificus (Bacillus), 815 otncolare (Bacillus), 815 otricolare (Endosporus), 815 ottolenghii (Bacillus), 803 ottolenghii (Clostridium) , 803 ottolenghii (Endosporus) , 804 ovalare (Clostridium), 796 ovalaris (Fleet ridi um) , 796 ovale (Bacterium) , 441, 683 ovale (Flavobacterium) , 441 ovale (Saccharobacterium), 623 oralis (Bacillus), 97, 441, 683 ovalis (Bacterium), 441 ovalis (Micrococcus), 270, 325, 326 ovalis (Pseud omonas), 97, 699 ovalis (Streptococcus), 325 ovalisporus (Myxococcus), 1043, 1044 ovata (Pasteurella) , 572 ovatum (Bacterium), 683 ovatum (Panhistophyton) , 254 ovatus (Bacteroides), 572, 577 ovatus (Micrococcus), 254 ovatus m,inutissimus (Bacillus), 683 ovatus mimdissimus (Bacterium) , 683 ovi (Pseudomonas) , 149 ovicola (Pseudomonas) , 149 oviforme (Bacterium), 353 oviformis (Bacteroides), 353 oviformis (Coccobacillus) , 353 orina (Ehrlichia), 1097 ovina (Rickettsia), 1097 ovina (Spirochaeta), 1068 ovina (Spiroschaudinnia) , 1068 ovinum (Bacterium,), 554 ovinum (Treponema) , 1068 ouzs (Corynebacterium), 62, 389, 694 ovis (Eperythrozoon) , 1112 or/s (Hemophilus), 589 oris (Listerella) , 409 ovzs (Micrococcus), 267 ovis (Spirillum.), 1068 ovis (Spirochaeta), 1068 oris (Spironema) , 1068 oris (Streptococcus), 342 ovis (Tortor), 1278 oris (Treponema), 1068 oviseptica (Pasteurella), 554 ovitoxicus (Bacillus), 790 ovitoxicus (Clostridium), 790, 826 ovitoxicus (Welchia), 790 ovoaethylicus (Bacillus), 720 oxalaticus (Bacillus), 714, 750 oxaliferum (Achromatium), 996, 997, 998, 999 oxydans (Acetobacter) , 184 oxydans (Bacillus), 182, 184 oxydans (Bacterium), 184 oxygenes (Bacterium), 544 oxygenes (Eberthella) , 544 oxygenes (Shigella), 544 oxylacticus (Bacillus), 664, 750 oxylacticus (Bacterium) , 750 oxyphila (Eberthella) , 534 oxxjphilum (Bacterium), 534 oxytocum (Aerobacter) , 456 oxytocum (Bacterium) , 456 oxytocus (Bacillus) , 456 oxytocus (Escherichia), io6 oxytocus perniciosus (Bacillus), 456 oxytocus perniciosus (Bacterium), 456 ozaenae (Bacillus), 459, 658 ozaenae (Bacterium), 459 ozaenae (Klebsiella), 459 ozenae (Encapsulata) , 459 pabuli (Bacillus), 750 pabuli (Plocamobacterium) , 693 pabuli acidi II (Bacillus) , 356, 693 pabuliacidi (Lactobacillus), 356 pachelabrae (Cristispira), 1056 pachyrhizi (Marmor), 1188 Pacinia, 192, 349, 763 paglianii (Bacillus), 656, 804 paglianii (Clostridium), 804 paglianii (Endosporus), 804 paleopneumoniae (Diplococcus), 309 pallens (Bacterium) , Q83 pallens (Micrococcus), 270 pallens (Streptococcus), 342 1477 pall-para INDEX OF NAMES OF GENERA AND SPECIES pallescens (Bacillus), 664 pallescens (Bacterium) , 683 pallescens (Pseudomonas) , 149, 699 palleum (Endosoma), 859 pallida (Spirochaeta) , 1071 pallida (Ulvina), 683 pallida var. cuniculi (Spirochaeta), 1073 pallidior (Bacterium) , 683 pallidolimbatus (Marmor) , 1197 pallidula (Spirochaeta), 1071 pallidulum (Treponema) , 1071 pallidum (Bacterium) , 683 pallidum (Microspironema) , 1071 pallidum (Spironema), 1071 pallidum (Treponema), 1071, 1072, 1073 pallidum \SiV. cunicidi (Treponema), 1073 pallidus (Bacillus), 664, 750 pallidus (Micrococcus) , 270 pallidus (Streptococcus), 342 palludosum (Plectridium) , 727 Palmula, 34, 763 paludis (Bacillus), 790, 826 paludis (Welchia), 790 paludosa (Sarcina) , 293 paludosum (Bacterium), 761 palumbarium (Bacterium), 552 palustris (Bacillus), 750 palustris (Hillhousia) , 998, 999 palustris (Rhodobacillus) , 750, 863 palustris (Rhodomonas) , 863 palustris (Rhodopseudomonas), 860, 863, 864, 865, 866 palustris var. gelaticus (Bacillus), 750 pammelli (Bacillus), 745 panacis (Pseudomonas), 130 panama (Salmonella), 519 panaxi (Bacterium), 130 panaxi (Phytomonas) , 130 pandora (Bacillus), 720 panginensis (Actinomyces), 921 panginensis (Nocardia), 921 panici (Bacterium), 169 panici (Phytomonas), 169 panici (Pseudomonas), 169 panici (Xanthomonas) , 169 panici -miliacei (Bacterium), 143 panici-miliacei (Phytomonas) , 143 panici-miliacei (Pseudomonas), 143 panificans (Bacillus), 664 pams (Bacillus), 710, 711 panis (Bacterium) , 760 panis (Lactobacillus), 358 panis acidi (Lactobacillus), 363 panis fermentati (Bacillus), 358 panja (Actinomyces), 966 panjae (Streptomyces) , 966 pannosus (Bacillus), 664 pannosus (Micrococcus) , 270 pansinii (Bacillus), 664, 667 pansinii (Bacterium) , 718 pansinii (Pseudomonas) , 149 pantotropha (Hydrogenomonas) , 77 pantotrophus (Bacillus) , 77 papavericola (Bacterium) , 164 papavericola (Phytomonas), 164 papavericola (Xanthomonas) , 164 papaveris (Bacillus), 474 papaveris (Erwinia), 474 papayae (Bacillus) , 478 papayas (Erwinia), 478 papillare (Bacterium) , 683 papillare (Photobacterium) , 637 papillaris (Microspira) , 637 pappulum (Paraplectrum) , 816 pappidus (Bacillus) , 815 papuana (Salmonella), 512 papulans (Bacterium) , 124 papidans (Phytomonas), 124 papulans (Pseudomonas), 124 paraabortus (Brucella), 563 para-aertrycke (Bacillus), 502 paraalvei (Bacillus), 723 paraamericanus (Proteus) , 491 para-asiatica (Salmonella), 532 para-asiaticus (Bacillus), 532 parabifermentans (Clostridium), 796 parabijermentans sporogenes (Bacillus ) , 796 parabifidus (Lactobacillus), 354 parabotulinum (Clostridium), 778, 779, 784 parabotulinum equi (Clostridium), 779 parabotulinum Types A and B (Clostrid- ium), 119, lS^,n^ 1478 INDEX OF NAMES OF GENERA AND SPECIES para-para parabotulinus (Bacillus), 779, 780 parabotidinus hovis (Clostridium), 77 parabutyricus (Bacillus), 813, 816 paracitrovorus (Streptococcus), 347 para-coagulans (Bacillus) , 532 para-coagulans (Sabnonella) , 532 Paracloster, 7, 705 paracoli (Bacterium), 460 Paracolobactrum, 460, 461, 489 para-colon (Bacillus), 460 para-colon (Salmonella), 460 paradifflue7is (Bacillus), 491 paradiffluens (Proteus), 491 paradoxa (Escherichia), 453 paradoxus (Bacillus), 683 paradoxus (Bacterium) , 683 paradoxus (Colibacillus), 453 paradysenteriae (Bacillus), 537, 538 paradyscntcriae (Bacterium) , 537 paradysenteriae (Eberthella) , 537 paradysenteriae (Shigella). 537, 538, 539, 540, 543 paradysenteriae Type Manchester (Shi- gella), 539 paradysenteriae (Type Xewcastle) (Shi- gella), 538 paradysenteriae var. sonnei (Shigella), 540 paradysenteriae X (Bacillus), 536 paraenterica (Enter aides) , 452 paraenterica (Escherichia), 453 paraentericus (Bacillus), 453 paraexilis (Bacillus), 352 paraffinae (Micrococcus) , 270 paraffinae (Nocardia), 901 paraffinae (Proactinomyces) , 901 para-gruenthali (Bacillus), 451 paragruenthali (Escherichia), 451 paraguayensis (Proactinomyces), 923 parainfluenzae (Hemophilus), 585, 586 paralacticus (Streptococcus), 324 parallela (Aphanothece) , 872 parallela (Pediochloris) , 872 parallelum (Pelodictyon), 872 parallclus (Bacillus), 664 paraluis (Spirochaeta) , 1073 paraluis-cuniculi (Spirochaeta), 1073 paralytica (Klebsiella) , 459 paralyticans (Bacillus), 405 paralyticans (Corynebacterium), 405 paramecii (Miillerina), 1122 paramelitensis (Brucella), 563 paramelitensis (Micrococcus), 563 paramorganii (Proteus), 491 paraoxytocum (Aerobacter) , 456 para-pertussis (Bacillus), 587 Paraplectrum, 7, 33, 763 paraputrificum (Clostridium), 793 paraputrificum (Plectridium) , 793 paraputrificus (Bacillus), 793 ■paraputrificus (Tissierillus), 793 paraputrificus coli (Bacillus), 793 parasarcophysematos (Bacillus), 775 parasitica (Leptothrix) , 366 parasiticum (Bacterium) , 366 parasiticum (Polyangium) , 1032 Paraspir ilium, 9 parasporogenes (Bacillus), 784 parasporogenes (Clostridium), 784 parasuis (Brucella) , 563 paratuberculosis (Mycobacterium), 881 paratyphi (Bacillus), 501 paratyphi (Bacterium), 501 paratyphi (Salmonella), 495, 501 paratyphi abortus ovis (Bacillus), 506 paratyphi alcaligenes (Bacillus), 501 paratyphi alvei (Bacillus) , 532 paratyphi A (Sabnonella) , 493, 501 paratyphi B (Salmonella), 493, 501 paratyphi C (Salmonella), 493, 507 paratyphi Typus A (Bacterium), 501 paratyphi, Typus B (Bacterium) , 501 paratyphosum A (Bacterium), 501 paratyphosum B (Bacterium), 501 paratyphosus (Bacillus), 501 paratyphosus A (Bacillus), 501 paratyphosus B (Bacillus), 501, 509 paratyphosus ^i (Bacillus), 507 paratyphosus B , Arkansas type (Bacillus), 508 paratyphosus B, Binns type (Bacillus), 503 paratyphosus B, Mutton type (Bacillus) 502 1479 para-pavo INDEX OF NAMES OF GENERA AND SPECIES paratyphosus B, Newport type {Bacillus), 513 paratyphosus B, Reading type (Bacillus,) 504 paratyphosus B, Stanley type (Bacillus), 503 paratyphosus C (Bacillus), 507, 509 paratyphosus C (Salmonella), 507, 509 paraviscosum (Bacterium) , 683 parcifermentans (Lactobacterium) , 364 parkeri (Borrelia), 1064 parkeri (Spirochaeta) , 1064 parotitidis (Micrococcus), 270 paroiitidis (Spirochaeta) , 1074 partum (Clostridium), 821 parva (Cornilia), 822 parva (Micromonospora) , 979 parva (Nocardia), 939 parvula (Cristispira) , 1056 parvula (Veillonella), 302, 303, 304 parvula var. branhamii (Veillonella), 303 parvula var. minima (Veillonella), 303 parvula var. thomsonii (Veillonella), 303 parvulum (Bacterium), 642 parvulum (Corynebacterium), 408 parvulus (Micrococcus), 302 parvulus (Staphylococcus), 302, 345 parvulus (Streptococcus), 331, 342 parvulus (Streptostaphylococcus) , 345 parvulus non liquefaciens (Streptococcus) , 331 parvum (Bacterium) , 822 parvum (Corynebacterium) , 388, 405 parvum (E ubacterium) , 368 parvum (Spirillum), 203, 701 parvum (Spirochaeta), 1075 parvum (Treponema) , 1075 parvum infectiosum (Corynebacterium), 405 parvus (Actinomyces) , 939 parvus (Bacillus), 544, 712 parvus (Jodococcus) , 270, 695 parvus (Micrococcus), 270 parvus (Phagus), 1131 parvus (Rhodovibrio) , 863 parvus (Streptomyces) , 939 parvus (Vibrio), 203 parvus liquefaciens (Bacillus), 404 parvus ovatus (Bacillus), 548 passeti (Staphylococcus), 256 passiflorae (Marmor), 1193 passiflorae (Phytomonas) , 138 passiflorae (Pseudomonas), 138 Pasteurella, 17, 21, 22, 26, 32, 37, 42, 546,551, 577, 1290 pasteuri (Cornilia), 115 pasteuri (Micrococcus) , 270, 306 Pasteuria, 35, 836 pasteuriana (Ulvina), 692 pasteurianum (Acetobacter), 61, 180, 182, 692 pasteurianum (Bacillus) , 692 pasteurianum (Bacterium), 182 pasteurianum (Clostridium), 772, 824 pasteurianum (Mycoderma), 182 pasteurianum (Rhyzobium), 225 pasteurianus (Bacillus) , 772 pasteurii (Bacillus), 729, 744 pasteurii (Urobacillus), 729 pasteurii (Vibrio), llA pastinator (Achromobacter) , 628 pastorianum (Clostridium) , 772 pastoriamnn (Lactobacterium), 360 pastorianum (Plocamobacterium) , 695 pastorianus (Bacillus) , 359, 772 pastorianus (Lactobacillus), 359, 695 pastorianus (Saccharobacillus) , 359 pastorianus (Streptococcus), 337, 342 pastorianus var. berolinensis (Saccharo- bacillus), 360 patelliforme (Bacterium) , 683 pater if ortne (Bacterium) , 683 pathogenicum (Photobacterium) , 635 paucicutis (Bacillus), 750 paullulus (Bacillus), 664 pauloensis (Eberthella) , 534 pauloensis (Escherichia) , 453 pauloensis (Salmonella), 532 Paulosarcina, 13, 285 paurometabolum (Corynebacterium), 392, pavlovskii (Bartonella), 1108 pavlovskii (Haemobartonella), 1108 pavonacea (Pseudomonas), 91 1480 INDEX OF NAMES OF GENERA AND SPECIES pavo-perf pavoninus (Bacillus) , 233 pavonis (Treponema), 1075 pecoris (Hostis), 1239 pectinis (Cristispira), 1056 Pectinobacter, 763 pectinophorae (Bacillus), 664 pectinovorum (Aerobacter), 456 pectinovorum (Bacillus), 822 pectinovorinn (Clostridium), 771, 822 pectinovorum (Granulobacter) , 771, 822 pectinovorum (Plectridiu?n), 771, 822, 824 pectinovorum liquefaciens (Plectridixwi) , 823 pectinovorus (Bacillus), 822 Pectobacillus, 8, 763 Pectobacterium, 464 pectocutis (Bacillus) , 750 pediculata (Nevskia), 830 pediculatum (Bacterium), 830 pediculatus (Chondromyces), 1038 pediculi (Bacillus) , 664 pediculi (Rickettsia), 1094, 1095, 1096, 1097 Pediochloris, 870 Pediococcus, 7, 25, 235, 249, 844 Pedioplana, 235, 250 pelagia (Bacillus), 635 pelagia (Bacterium), 635 pelagia (Sarcina), 701 pelagicus (Bacillus), 750 pelamidis (Spirochaeta) , 1068 pelamidis (Spironema), 1068 pelargoni (Bacterium), 160 pelargoni (Pseudo7no7ias), 160 pelargonii (Marmor), 1199 pelargonii (Phytomonas) , 160 pelargonii (Xanthomonas), 160, 167 pelletieri (Actinomyces), 960 pelletieri (Discomyces) , 960 pelletieri (Micrococcus), 960 pelletieri (Nocardia), 960 pelletieri (Oospora), 960 pelletieri (Streptomyces), 960 pellicidosa (Pseudomonas) , 149 pellucida (Beggiatoa) , 992 pellucida (Pseudomonas), 149 pellucidmn (Achromobacter) , 145 pellucidum (Halibacterium) , 653 pellucidus (Bacillus), 664, 750 pellucidus (Micrococcus), 270 Pelochromatium, 859 Pelodictyon, 870 pelogenes (Actinomyces), 972 Pelogloea, 870 pelomyxae (Cladothrix) , 1123 pelurida var. virginiana (Pseudomonas) , 174 pemphigi (Micrococcus), 270 pemphigi acuti (Diplococcus) , 270 pemphigi contagiosa (Micrococcus), 270 pemphigi neonatorum (Micrococcus), 271 pemphigi neonatorum (Staphylococcus), 271 pendens (Rhodothece),855 pendunculatus (Bacillus) , 741 penicillatus (Bacillus), 816 penortha (Spirochaeta) , 583, 917, 1074 pensacola (Salmonella) , 518 pentoaceticum (Plocamobacterium) , 695 pentoaceticus (Lactobacillus), 358, 695 pentoaceticus var. magnus (Lacto- bacillus), 358 pentosaceum (Propionibacterium), 378, 379 pentosaceus (Pediococcus), 250 pentosus (Lactobacillus) , 357 pepo (Bacillus), 730 Peptoclostridium, 30, 763 Peptococcus, 29, 30 peptogenes (Bacillus), 751 peptogenes (Bacterium) , 751 peptonans (Bacillus), 751 peptonificans (Bacillus), 751 Peptonococcus, 235 Peptostreptococcus , 30, 31, 312 percitreus (Micrococcus) , 271 percolans (Vibrio), 201 perekropovi (Eperythrozoon) , 1113 per exile (Treponema), 1075 perexilis (Spirochaeta), 1076 perfectomarinus (Pseudomonas) , 699 perflava (Neisseria), 298 perflavus (Micrococcus), 271 1481 peif-pete INDEX OF NAMES OF GENERA AND SPECIES perfoetens (Bacterium), 576 perfoetens (Bacteroides) , 576 perfoetens {Ristella), 576 perforans (Spirochaeta) , 1068 perfringens (Bacillus), 790, 815, 826 perfringens (Clostridium), 789, 790, 817, 818, 826 perfringens Type A (Clostridium), 789 perfringens Type B (Clostridium), 790 perfringens Type C (Clostridium), 790 perfringens Type D (Clostridium), 790 perfringens (Welchia), 790 perfringens var. anaerogenes (Clostrid- ium), 791, 826 perfringens var. e!7cns (Clostridium), 790 perfringens var. c^e/is (If'e/c/u'u), 790 perfringens var. zoodysenteriae (Welchia), 791 pericarditis (Paste urella), 554 pcricoma (Vibrio), 205 perimastrix (Vibrio), 703 periphyta (Pseudomonas) , 699 periplanetac (Corynebacterium) , 405 periplanetae (Spirochaeta) , 1069 periplanetac var. americana (Coryne- bacterium), i05 periplaneticum (Spirillum), 217 peripneumoniae (Borrelomyces), 1291 peripneumoniae (Mycoplasma), 1291 peripneumoniae bovis (Asleromyces) , 1291 peripneumoniae bovis contagiosae (Mi- cromyces) , 1291 peritonitidis (Streptococcus), 317 peritonitidis equi (Streptococcus), 317 peritonitis (Spherophorus), 579 peritlomaticum (Bacterium), 761 perlibratus (Bacillus), 664 perlucidulus (Bacillus), 751 perlurida (Cellulomonas), 174 perlurida (Pseudomonas), 174 perniciosus (Micrococcus), 342 perniciosus (Pediococcus) , 250 perniciosus (Streptococcus), 342 perniciosus psittacorum (Streptococcus), 342 perolens (Achromobacter) , 425 peromysci (Grahamella) , 1111 peromysci (Haemobartonella), 1106 peromysci var. maniculati (Grahamella), nil peromysci var. maniculati (Haemobar- tonella), 1106 peromelia (Bacillus), 816 peroxydans (Acetobacter), 189 perroncitl (Bacillus), 664 Perroncitoa, 312 persica (Borrelia) , 1069 persica (Spirochaeta) , 1069 persicae (Chlorogenus) , 1148, 1152 persicae (Flavimacula) , 1196 persicae (Marmor), 1196, 1202 persicae var. micropersica (Chlorogenus) , 1149 persicae var. vidgaris (Chlorogenus), 1149 persicina (Palmella), 848 persicina (Sarcina), 293 'persicum (Treponema), 1069 persicus (Micrococcus), 271 pertenue (Spironema) , 1071 pertenue (Treponema), 1071 pertenuis (Spirochaeta), 1071 pertussis (Bacillus), 586, 737 pertussis (Hemophilus), 586, 587, 589 pertussis eppendorf (Bacillus) , 589 peruviana (Bartonella), 1101 pestifer (Achromobacter) , 425 pesiifer (Bacillus), 425 pestifer (Bacterium) , 425 pesizs (Bacillus), 549 pestis (Bacterium) , 549 pestis (Eucystia), 549 pestis (Pasteurella), 549, 703 pestis bubonicae (Bacillus), 549 pestis bubonicae (Bacterium), 549 pestis-caviae (Bacillus), 502 pestis-caviae (Pasteurella) , 502 petasites (Bacillus), 714 petasitis (Bacterium) , 142 petasitis (Phytomonas) , 142 petasitis (Pseudomonas), 142 petechialis (Micrococcus) , 271 petersii (Bacillus), 664 1482 INDEX OF NAMES OF GENERA AND SPECIES pete-phos petersii {Bacterium), 683 peterssonii (Propionibacteriuni), 376 petilus {Micrococcus) , 271 ■petiolatus {Bacillus) , 751 ■petit {Spirochaeta) , 215 petrolei {Micrococcus) , 271 petroselini {Bacillus) , 715 petroselini {Bacterium), 715 pettiti {Leptospira), 1079 pettiti {Spirochaeta), 215, 1079 pettiti {Treponema), 1079 P/a# {Bacillus), 539 pfaffii {Bacterium), 539 p/aj^i {Eberthella) , 5ZQ pfaffii (Shigella), 539 Pfeifferella,.9, 21, 22, 28, 554 pfeifferi {Coccobacillus), 585 pjeifferi {Encapsulatus) , 459 pfiegmones-emphrjsematosae (Bacillus) , 790 pflegviones-emphysematosae {Clostridium) , 790 pflilgeri {Arthrobacterium) , 635 pfliigeri {Bacterium,) , 635 pfliigeri {Micrococcus), 635 pfliigeri {Photobacterium) , 635 Phacelium, 13, 285 phaeochroviogenus {Actinomyces), 943 phaeochromogenus (Streptomyces), 943 Phaeomonas, 29, 30 Phaeospirillum, 29, 30, 866 phagedenis {Borrelia), 1064 phagedenis {Spirochaeta), 1064 phagedenis (Spironema) , 1064 phagedenis {Spiroschaudinnia) , 1064 phagedenis {Treponema), 1064 Phagus, 1128 pharyngis {Diplococcus) , 299 pharyngis (Micrococcus), 696 pharyngis (Neisseria), 299, 696 pharyngis (Staphylococcus), 282 pharyngis cinerea (Neisseria), 298 pharyngis cinereus (Micrococcus), 298 301 pharyngis communis (Diplococcus), 298 pharnygis flavus I (Diplococcus), 298 pharyngis flavus II (Diplococcus), 299 pharyngis flavus I (Micrococcus) , 299 pharyngis flavus II (Micrococcus), 299 pharyngis-sicci (Neisseria) , 298 pharyngis siccus (Diplococcus), 298 pharyngis siccus (Micrococcus), 298 phaseoli (Bacillus), 160, 751 phaseoli (Bacterium), 160 phaseoli (Phytomonas) , 160 phaseoli (P seudomonas) , 160 phaseoli (Marmor), 1168, 1179 phaseoli (Rhizobium), 225, 226 phaseoli (Xanthomonas) , 160, 161, 1134, 1136 phaseoli var. fuscans (Bacterium), 161 phaseoli var. ftiscans (Phytomonas) , 161 phaseoli var. fuscans (P seudomonas) , 161 phaseoli var. /wsca?js (Xanthomonas) , 161 phaseoli var. sojense (Bacterium), 161 phaseoli var. sojense (Phytomonas), 161 phaseoli Y&T.sojensis (Xanthomonas), 161 phaseolicola (Pseudomonas), 118 phasiani (Bacillus), 520 phasiani septicus (Bacillus), 520 phasiani septicus (Bacterium), 520 phasianicida (Bacillus), 552 phasianicida (Bacterium), 552 phasianidarum mobile (Bacterium), 552 phenanthrenicus bakiensis (Bacillus) , 664 phenanthrenicus guricus (Bacillus), 665 phenologenes (Bacillus), 665 phenolphilos (Bacillus), 751 phenotolerans (Actinomyces), 917 phlebotomi (Spirochaeta), 1074 phlegmones emphysematosae (Bacillus), 789, 790, 826 phlei (Mycobacterium), 881, 887, 889, 890 p/iZcz (Sclerothrix) , 889 p/iZei perrugosum (Mycobacterium), 890 p/iZei planum (Mycobacterium) , 890 Phleobacterium, 20, 23 pholas (Bacterium), 635 phormicola (Bacterium), 166 phormicola (Phytomonas), 166 phormicola (Xanthomonas), 166 phosphorescens (Bacillus), 111 1483 phos-pinn INDEX OP NAMES OF GENERA AND SPECIES phosphorescens (Bacterium), 111, 633, 635, 636 phosphorescens (Micrococcus), 633 phosphorescens (Microspira) , 700 phosphorescens (Pasteurella), 699 phosphorescens (Photobacteriimi), 633, 635, 636 phosphorescens (Pseudomonas) , 111, 699 phosphorescens (Spirillum), 633 phosphorescens (Vibrio), 633, 702 phosphorescens gelidus (Bacillus), 636 phosphorescens gelidus (Bacterium), 635 phosphorescens giardi (Bacillus), 635 phosphorescens giardi (Bacterium), 635 phosphorescens indicus (Bacillus), 699 phosphorescens indicus (Bacterium), 699 phosphorescens indigenus (Bacillus), 633 phosphorescens indigenus (Bacterium), 633, 636 phosphorescens pjlugeri (Bacterium), 635 phosphoreum (Achromobacter) , 634 phosphoreum (Bacterium), 633, 635, 636, 637 phosphoreum (Microspira), 636 phosphoreum (Phoiobacter), 633 phosphoreum (Phoiobacter ium) , 633 phosphoreus (Bacillus) , 633, 634 phosphoreus (Micrococcus), 633, 635 phosphoreus (Streptococcus), 633 phosphoricum (Achromobacter) , 634 phosphoricus (Bacillus), 634 Phoiobacter, 636 Phoiobacter ium, 12, 14, 192, 636 phoiometricum (Bacterium), 683 photometricum (Rhodospirillum) , 867 photometricus (Bacillus), 683 Photomonas, 11, 636 Photospir ilium, 636 photuris (Proteus), 491 Phragmidiothrix, 6, 12, 17, 19, 987 phyllotidis (Grahamella) , 1111 physiculus (Micrococcus), 636 Phyiomonas, 31, 32, 150 Phytomyxa, 223 phytophthora (Erwinia), 469, 470 phytophthorum (Bacterium), 470 phytophthorum (Pectobacterium) , 470 phytophthorus (Bacillus), 469, 470 phytophthorus (Micrococcus) , 342 phytophthorus (Streptococcus), 342 phytoplanktis (Vibrio), 703 picrogenes (Bacillus), 751 picrum (Achromobacter) , 622 pictor (Treponema), 1072 pictorum (Achromobacter), 177 pictorum (Pseudomonas), 177 pierantonii (Bacillus), 202 pierantonii (Cocco-bacillus), 112 pierantonii (Micrococcus), 112 pierantonii (Pseudomonas), 112 pierantonii (Vibrio), 202 pieridis (Micrococcus), 271 pieris (Borrelina), 1227 pieris (Diplobacillus) , 690 pieris (Diplococcus) , 336 pieris (Vibrio), 703 pieris agilis (Bacillus) , 665 pieris fluorescens (Bacillus) , 665 pieris liquefaciens (Bacillus) , 665 pieris liquefaciens a (Bacillus), 665 pieris liquefaciens /3 (Bacillus) , 665 pieris liquefaciens (Bacterium) , 665 pieris non-Uquefaciens a (Bacillus) , 665 pieris non-Uquefaciens ^ (Bacillus) , 665 piesmae (Savoia), 1221 piima (Streptococcus) ,Si2 pijperi (Actinomyces), 921 pijperi (Discomyces) , 921 pijperi (Nocardia) , 921 pikowskyi (Achromobacter), 425 pikowskyi (Micrococcus), 271 piliformis (Bacillus), 366, 751 piliformis (Micrococcus), 271 piltonensis (Micrococcus), 271 piluliformans (Bacillus), 684 piluliformans (Bacterium) , 683 pmi (Bacterium), 640 pwi (Pseudomonas), 640 pinnae (Cristispira), 1055 pinnae (Spirochaete) , 1055 pinnatum (Achromobacter) , 425 pinnatus (Bacillus), 425 pinnatus (Bacterium), 426 1484 INDEX OF NAMES OF GENERA AND SPECIES pino-plic pinoyi (Actinomyces), 921 pinoyi {Nocardia), 921 pintae {Treponema), 1072 pipientis (Wolbachia), 1098 pipistrelli (Grahamella), 1111 pipistrelli (Grahamia), 1111 piriforme {Corynebacterium) , 694 piscatora (Shigella), 544 piscatonim (Bacterium), 483 piscatorum (Serratia), 483 piscatorus (Bacillus) , 483 pisces (Rickettsia), 1097 piscicidus (Bacillus), 751 piscicidus agilis (Bacillus) , 751 piscicidus agilis (Bacterium) , 751 piscicidus nobilis (Bacillus) , 751 piscicidus versicolor (Bacillus), 491 piscicidus versicolor (Proteus), 491 l)iscium (Mj'cobacterium), 883, 884, 886, 887 piscium (Vibrio), 197 piscova (Pseudomonas), 700 pisi (Bacillus), 751 pisi (Bacterium), 119 ]>isi (Marmor), 1180 pisi (Phytomonas), 119 pisi (Pseudomonas), 119 pistiense (Thiospirillum) , 853 pitheci (Spirillum) , 1064 pitheci (Spirochaeta) , 1064 pitheci (Spironema) , 1064 pituitans (Bacterium), 761 pituitoparus (Micrococcus), 271 pituitoparus (Karphococcus) (Car coccus), 271 pituitosum (Bacterium), 425, 684 pituitosum (Propionibacterium) , 379 pityocampae (Bacterium), 684 pitycampae c (Streptococcus) , 342 pitycampac j3 (Streptococcus), 342 placoides (Cladothrix), 974 placoides (Leptotrichia), 974 placoides alba (Leptothrix), 974 plagarum (Injlabilis), 823 plagarum-belli (Diplococcus), 310 Planococcus, 7, 13, 15, 235 Planomerista, 235 Planosarcina, 7, 15, 285 Planostreptococcus, 15, 312 plantaginis (Phytomonas), 161 plantaginis (Xanthomonas), 161 plantarum (Lactobacillus), 356, 357, 363, 695 plantarum (Streptobacterium), 356 plantarum var. rtidensis (Lactobacillus) , 357 platus (Bacillus), 723 platychoma (Bacillus), 751 plauti (Fusocillus), 581 plauti-vincenti (Fusobacterium), 581, 1063 plauti-vincenti (Fusiformis), 581 plant -vincenti (S pirochaete) , 1063 plebia (Escherichia) , 451 885, plebeius (Bacillus) , i5l Plectridium, 7, 13, 33, 763 Plectrillium, 7 Plectrinium, 7 plehniae (Pseudomonas) , 149 Plennobacterium, 705 pleofructi (Bacillus), 346 pleofructi (Leuconostoc) , 346 pleomorpha (Pseudomonas), 700 pleomorphus (Bacillus), 665 pleomorphus (Streptococcus), 342 pleuriticus (Discomyces) , 915 pleuriticus (Leptothrix), 915 pleuriticus canis familiaris (Actinomy- ces), 915 Pleuropneumonia, 1289 />^'«- pleuropneumoniae (Bacterium), 684 pleuropneumoniae (Bovimyces), 1291 plexiformis (Bacillus), 751 plicata (Pseudomonas), 149 plicatile (Spirillum), 1052 plicatilis (Ehrenbergia), 1052 plicatilis (Spirochaeta), 1052, 1053 plicatilis (Spirulina), 1052 plicatilis eurystrepta (Spirochaeta), 1052 plicatilis inarina (Spirochaeta) , 1052 plicatilis plicatilis (Spirochaeta), 1052 plicativum (Bacterium) , 684, 761 plicatum (Acetobacter), 186 1485 plic-poly INDEX OF NAMES OP GENERA AND SPECIES plicatum (Bacterium), 761 plicatum (Flavobacterium) , 441 plicatus (Bacillus), 149, 441, 665, 684, 747, 751 Plocamobacterium, 18, 25, 27, 349, 381, 400 plumbeus (Bacillus), 665 plumosum (Corynebacterium) , 405 plumosum (Mycobacterium) , 405 plumosus (Micrococcus) , 271 plurichromogenes (Actinomyces), 897 pluricolor (Actinomyces) , 973 pluricolor (Nocardia), 973 pluricolor (Streptothrix), 973 pluricolor diffundens (Actinomyces) , 973 pluricromogena (Streptothrix), 897 pluriseptica (Pasteurella), 547 plurisepticus (Bacillus), 546 pluton (Bacillus), 724 pluton (Diplococcus) , 724 plymouthensis (Bacillus) , 482 plymouthensis (Erythrobacillus) , 482 plymuthicum (Bacterium) , 482 plymuthicum (Serratia), 481, 482, 483, 484, 485 Pneumobacillus, 458 Pneumococcus , 306 pneumo-enteritidis murium (Bacillus), 665 pneumoniae (Bacillus), 458, 652 pneumoniae (Bacterium), 307, 458 pneumoniae (Diplococcus), 43, 45, 306, 307, 308, 309 pneumoniae. Type 3 (Diplococcus), 308 pneumoniae (Encapsulatus) , 458 pneumoniae (Hyalococcus), 458 pneumoniae (Klebsiella), 458 pneumoniae (Miyagawanella), 1118, 1119 pneumoniae (Pneumococcus), 307 pneumoniae (Proteus), 458 pneumoniae (Streptococcus), 307 pneumoniae caprae (Bacillus), 553 pneumoniae crouposae (Micrococcus) , 307 pneumoniae equi (Bacillus), 553 pneumonicum (Bacterium), 458, 675 pneumonicus (Bacillus), 647 pneumonicus agilis (Bacillus), 647 pneumonicus agilis (Bacterium), 647 pneumonicus liquefaciens (Bacillus), 675 pneumonicus liquefaciens (Bacterium), 675 pneumonic crouposae (Bacterium), 458 pneumopecurium (Bacterium) , 684 pneumosepticum (Bacterium), 684 pneumosepticus (Bacillus) , 665, 684 pneumosepticus (Bacterium) , 665 pneumosimilis (Streptococcus) , 342 pneumosintes (Bacillus), 595 pneumosintes (Bacterium), 595 pneumosintes (Dialister), 595 Podangium, 1009, 1034 podovis (Treponema) , 1074 poeciloides (Bacillus), 368, 369 poeciloides (Eubacterium) , 368, 369 poelsii (Bacillus), 665 poinsettiae (Corynebacterium), 398, 399 poinsettiae (Phytomonas), 399 pollacii (Bacillus), 751 pollachii (Spirochaeta) , 1069 pollachii (Spironema), 1069 Pollendera, 705 Polyangium, 5, 14, 17, 20, 24, 26, 1005, 1009, 1025 Polycephalum, 1036 polychromogenes (Actinomyces) , 897 polychromogenes (Bacillus), 233 polychromogenes (Nocardia), 897, 898 polychromogenes (Oospora), 897 polychromogenes (Proactinomyces) , 897 polychromogenes (Streptothrix), 897 polycolor (Bacterium), 126 polycolor (Phytomonas) , 126 polycolor (Pseudomonas), 89, 126 polycystus (Chondrococcus), 1045 polycystus (Myxococcus) , 1045 polydorae (Cristispira), 1056 polydorae (Cristispirella) , 1057 polyfermenticum (Clostridium), 772, 824 polygoni (Phytomonas), 140 polygoni (Pseudomonas), 140 polymorpha (Mima), 595 polymorpha var. oxidans, 595 polymorphum (Bacterium), 684 polymorphum (Fusobacterium), 582 polymorphum (Halibacterium) , 662 1486 INDEX OF NAMES OF GENEEA AND SPECIES poly-prod polyrnorphum (Rhizobium) , 224 polymorphum convulsivum (Bacterium), 590 polymorphus (Actinobacter) , 456 polymorphus (Bacillus), 684 polymorphus (Ftisiformis) , 582 polymorphus (Streptococcus) , 343 polymorphus (Vibrio), 205 polymorphus necroticans (Coccobacillus), 690 polymorphus necroticans (Micrococco- bacillus), 690 polymorphus var. peritriche (^^ibrio), 205 polymyxa (Aerobacillus), 720 polymyxa (Bacillus), 720, 722, 754, 818 polymyxa (Clostridium), 720 polymyxa (Granulobacter) , 720 polymyxa var. mucosum (Granulobacter) , 720 polymyxa var. ie/iax (Granulobacter) , 720 polypiformie (Cornilia), 816 polypiformis (Bacillus), 816, 818 polypus (Micrococcus), 272 polysclerotica (Spirochaeta) , 1065 polysiphoniae (Bacterium), 624 polysiphoniae (Flavobacterium) , 624 polyspira (Spirochaeta), 1074 polyspirum (Treponema), 1074 polyspora (Crenothrix), 983, 987 polyspora (Metabacterium) , 762 polytrophum (Nitrobacter) , 76 pomodoriferus (Bacillus), 665 pomona (Salmonella) , 529 poncei (Bacillus), 665 ponceti (Actinomyces), 921 ponceti (Nocardia), 921 ponceti (Oospora), 921 ponticus (Vibrio), 703 Pontothrix, 1002 poolensis (Actinomyces), 949 poolensis (Streptomyces), 949 poona (Salmonella), 527, 531 popilliae (Bacillus), 727 populi (Bacillus), 751 populi (Micrococcus) , 272 porcellorum (Micrococcus), 272 porci (Erysipelothrix) , 410 porri (Bacillus), 684 porrj (Bacterium), 684 portae (Spirillum), 1052 portuensis (Microspira) , 205 portuensis (Vibrio), 205 postumus (Bacillus), 816 potens (Acetobacter) , 692 potsdam (Salmonella), 511 potsdamensis (Salmonella), 511 praeacuta (Zuberella), 577 praeacutus (Coccobacillus), 577 praecox (Actinomyces) , 973 praefecundus (Actinomyces) , 973 praepollens (Bacillus), 665 praussnitzii (Bacillus), 718 praussnitzii (Bacteroides) , 577 preisz-nocardi (Corynebacterium) , 389 premens (Tarpeia), 1270 preputialis (Leptothrix), 366 Pretoria (Salmonella), 526 pretoriana (Nocardia), 900 pretorianus (Actinomyces), 900 prillieuxianus (Bacillus), 132 prima (Vibrio), 205 primarius (Phagus), 1132 primigenium (Archangium), 1018, 1019 primigenium (Polyangium), 1018 primigenium var. assurgens (Arch- angium), 1006, 1018 primulae (Marmor), 1201 primulae (Phytomonas) , 114 primulae (Pseudomonas), 114 primus (Robertsonillus), S23 primus fullest (Bacillus), 666 primus fullesi (Bacterium), 666 pringsheim (Bacillus) , 819 priztnitzi (Bacillus), 534 priztnitzi (Bacterium) , 534 priztnitzi (Eberthella) , dS-i priztnitzi (Eberthus), 534 Proactinomyces, 38, 875, 892 Proactinomyces sp., Helzer, 923 probatus (Bacillus), 729 prodeniae (Bacterium) , 684 prodigiosa (Micraloa) , 480 prodigiosa (Monas), 480 prodigiosa (Palmella), 480 1487 prod-pseu index of names of genera and species prodigiosa {Salmonella), 480 prodigiosum {Bacteridium), 480 prodigiosum {Bacterium) , 480 prodigiosum {Chromohacterium) , 480 prodigiosum {Dicrobactrum) , 480 prodigiosum (Liquidobacterium), 480 prodigiosus {Bacillus), 480 prodigiosus {Erythrobacillus), 480 prodigiosus {Micrococcus), 480 productus {Streptococcus), 343 profundus {Cladothrix) , 983 profusum {Bacterium) , 684 profusus {Bacillus), 684 progrediens {Micrococcus), 272 progrediens {Reglillus), 823 prolifer {Vibrio), 213 promissus {Bacillus) , 666 proper a {Thiospira), 702 Propionibacterium, 30, 31, 372 propionici {Micromonospora) , 980 Propionicoccus , 11 propionicum {Clostridium) , 821 propyl-butylicurn {Clostridium), 781, 825 Proshigella, 535 Protaminobacter, 32, 189 protea {Actinomyces), 972 protea {Microspira) , 197 protea {Pseudomonas) , 149 protea-fluorescens {Pseudomonas) , 89 proteamaculans {Bacterium), 169 proteamaculans {Phytomonas) , 169 proteamaculans {Pseudomonas), 169 proteamaculans (Xanthomonas) , 169 proteidis {Bacillus) . 666 proteiformis {Enterococcus), 325 proteiformis {Streptococcus) , 325 proteiformis liquefaciens {Enterococcus), 343 proteiformis var. liquefaciens {Strepto- coccus), 343 proteolyticum {Clostridium), 821 proteolyticum {Plocamobacterium) , 691 proteolyticus {Glycobacter) , 691 proteolyticus {Martellillus), 823 proteosimilis {Eberthella) , 534 protervus {Bacillus) , 666 proteum {Bacterium), 718 Proteus, 18, 21, 26, 31, 486, 489, 490, 491, 689, 691, 1086, 1088, 1089, 1091, 1092, 1093, 1103 proteus {Bacillus), 486 proteus {Bacterium), 691 proteus (Flavobacterium), 438 proteus {Oospora), 973 proteus {Streptococcus), 343 proteus {Streptothrix) , 973 proteus (Vibrio), 197, 202, 204, 205 proteus anindolgenes {Bacterium) , 487 proteus fluorescens {Bacillus) , 89 proteus fluorescens {Bacterium) , 698 proteus mirabilis {Bacillus), 488 proteus septicus {Bacillus), 686 proteus vulgaris {Bacillus), 487, 632 proteus-zenkeri {Bacillus) , 608 Protobios, 1128, 1129 protobios {Protobios), 1128 protozoides {Bacterium), 145 prowazeki sub-species